Erwin 2007 Caraboidea of the Western Hemisphere (Carabidae)

Asynthesis 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 occur-

rence, 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 indis-

pensable 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).

Cover: Cicindis horni Bruch

Pensoft Series Faunistica 66

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

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TRIBE ASDASDASD | 1

A TREATISE ON THE WESTERN

HEMISPHERE CARABOIDEA (COLEOPTERA)THEIR CLASSIFICATION, DISTRIBUTIONS, AND WAYS OF LIFE

VOLUME I (TRACHYPACHIDAE, CARABIDAE – NEBRIIFORMES 1)

Terry L. Erwin

2 | A TREATISE ON THE WESTERN HEMISPHERE CARABOIDEA. VOLUME I

This page intentionally left blank

TRIBE ASDASDASD | 3

SOFIA–MOSCOW

2007

A Treatise on the WesternHemisphere Caraboidea (Coleoptera)

Their classification, distributions,and ways of life

Volume I(Trachypachidae, Carabidae – Nebriiformes 1)

TERRY L. ERWIN


Pensoft PublishersGeo Milev Str. 13a, Sofia 1111, Bulgaria

Fax: [email protected]

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 2007ISBN 978-954-642-298-9

Pensoft Series Faunistica No 66ISSN 1312-0174

© PENSOFT Publishers

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system ortransmitted in any form by any means, electronic, mechanical, photocopying, recording or

otherwise, without the prior written permission of the copyright owner.

Printed in Bulgaria, April 2007

TRIBE ASDASDASD | 5

Dedicated To

Professor George Eugene BallUniversity of AlbertaEdmonton, Canada

GEORGE E. BALL

PICO DE NEBLINA, VENEZUELA, 1984SMITHSONIAN EXPEDITION

George likes books and little brooksthose hard-to-reach places in whichcarabid beetles dwell; landscapes

that make the muscles twitch n’ swellthat’s when George Ball

hears yon the Beckon Calltaking the student – he mentors! shapes!

the next generation’s itchto add yet another species to the Tell

TRIBE ASDASDASD | 7

Contents

Foreward 8

Introduction 9

Methods 10

Volume supplements and Database 13

Bibliography 13

Images 13

Acknowledgements 15

Arrangement of Western Hemisphere taxa: Caraboidea 16

Species accounts 17

Family TRACHYPACHIDAETribe Systolosomatini, Systolosoma 17Tribe Trachypachini, Trachypachus 18

Family CARABIDAETribe Pelophilini, Pelophila 20Tribe Nebriini, Leistus, Nebria, Nippononebria 22Tribe Notiokasiini, Notiokasis 51Tribe Opisthiini, Opisthius 52Tribe Notiophilini, Notiophilus 53Tribe Cicindini, Cicindis 61Tribe Omophronini, Omophron 63Tribe Loricerini, Loricera 69Tribe Carabini, Carabus, Calosoma, Callisthenes, Calopachys 72

Subtribe Ceroglossina, Ceroglossus 118Tribe Cychrini, Cychrus, Scaphinotus, Sphaeroderus 137Tribe Collyridini, Ctenostoma 171

General Bibliography for Caraboidea of the Western Hemisphere 207

Index 317

Plates 325


ForewardGoing on nearly 4 decades of admiration for my dear friend and still mentor, toProfessor George E. Ball, I dedicate these volumes, hard bound with carabid imagesand 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 aboutthem, mentoring, apprenticing, training, and teaching young carabid workers anddozens of other-taxa students, building important reference collections, and sharinghis time with anyone needing a carabid identification, their distributions, or generallife philosophy. Without him, and all those Carabidologists and generalist beetlecollectors he trained or influenced, much of what is included in these volumes wouldnot be available to the world of beetle enthusiasts and natural historians, indeed, tothe world in general.

TERRY L. ERWIN

National Museum of Natural HistorySmithsonian InstitutionWashington, DC

Introduction

Inspired by the book, A Natural History of the Ground Beetles (Coleoptera: Carabidae) ofAmerica North of Mexico (Larochelle, A. & Larivière, M.C. 2003), its utility and eleganceon 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 theLarochelle & Larivière contribution, I decided to migrate the information in myelectronic database of Western Hemisphere Caraboidea to the printed page. Thereare at present 9,242 species and subspecies currently recorded from the WesternHemisphere, therefore this Treatise will necessarily appear in six or seven volumes,with subsequent supplemental updates. The electronic version is updated on a sixmonth basis. These volumes will add to and update the species of North Americadocumented by Larochelle & Larivière (2003) with new information garnered fromdata in the large collections at NMNH, CAS, CMNH, UASM, and others, as well asmy personal field notes from 39 years of neotropical field work, thus extendingLarochelle & Larivière’s important contribution south of the U.S. - Méxican borderby treating species of northern México, the Neotropics and Neaustral Regions.Together, my volumes and that of Larochelle & Larivière can be considered as a setof handbooks on the ‘ways of life’ of Western Hemisphere carabid beetles. Larochelle& Larivière included information that I don’t cover, for example laboratory feedingand trapping methods of the North American species, while I include informationthey didn’t include, for example, historical nomenclature, distributions, altitudinalinformation, and data from specimens in major collections not heretofore published,as well as color images of adults and landscapes/habitats. We both also contributecomplementary bibiliographies. Nearly all carabid species need much more study beforewe 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 regardedas a starting point for further discoveries, many of which will be exciting and worthtelling again and again sometime, somewhere, over a camp fire. The “Tell” (Film: MadMax Beyond Thunderdome, 1985) began with us in the early evolution of humankind asoral 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 modeby which future generations will ‘tell,’ however, the written page stored amongstdispersed libraries remains, at least today, as the only reliably permanent continuum ofhuman knowledge.

INTRODUCTION | 9


Methods

For over three decades, data on Caraboidea of the Western Hemisphere have beenstored in various data management systems in my Tropical Forest Canopy Laboratoryat the National Museum of Natural History, Smithsonian Institution. In the early1990’s, these data were migrated to FileMakerPro and launched on the Departmentof Entomology website. Updates were made to the website, as reprints were receivedfrom colleagues, and at each new issue of the Zoological Record (Coleoptera). Plansare now being made to move this carabid database into SEQUEL, another databasemanagement system that is more secure on today’s internet, in anticipation of theSmithsonian’s planned Encyclopedia of Life Project. In addition, a version of theinformation 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 muchas possible. Organization of higher taxa follows that of Erwin (1985) with somemodifications based on recent publications. Within each Tribe, genera are arrangedalphabetically, 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 Patagonianregion. 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

Table 1. Abbreviations of states and provinces use in the species accounts.

METHODS | 11

AB AlbertaAG AguascalientesAI Aleutian IslandsAK AlaskaAL AlabamaAR ArkansasAZ ArizonaBC British ColumbiaBJ Baja CaliforniaCA CaliforniaCH ChihuahuaCL CoahuilaCM ColimaCO ColoradoCP CampecheCS ChiapasCT ConnecticutDC District of ColumbiaDE DelawareDF Distrito FederalDU DuragngoFL FloridaGA GeorgiaGJ GuanajuatoGO GuerreroHD HidalgoaIA IowaID IdahoIL IllinoisIN IndianaJA JaliscoKS KansasKY KentuckyLA LouisianaLB LabradorMA MassachusettsMB ManitobaMD MarylandME MaineMH MichoacanMI MichiganML MorelosMN MinnesotaMO MissouriMS MississippiMT MontanaMX México -NA Nayarit

NE NebraskaNC North CarolinaNB New BrunswickNF NewfoundlandNH New HampshireNJ New JerseyNK North DakotaNL Neuvo LeonNM New MexicoNS Nova ScotiaNT Northwest TerritoriesNV NevadaNY New YorkOA OaxacaOH OhioOK OklahomaON OntarioOR OregonPA PennsylvaniaPE Prince Edward IslandPM St. Pierre and MiquelonPQ QuebecPU PueblaQR Quintana RooQT QueretaroRI Rhode IslandSK SaskatchewanSC South CarolinaSD South DakotaSI SinaloaSL San Luis PotosiSO SonoraTA TabascoTL TlaxcalaTM TamaulipasTN TennesseeTX TexasUT UtahVA VirginiaVC VeracruzVT VermontWA WashingtonWS WisconsinWV West VirginiaWY WyomingYT Yukon TerritoryYC YucatanZA Zacatecas


An extended focus image or digital image of an exemplar adult of some speciesin each genus is included and where possible, an exemplar image is provided for eachgenus of a living beetle and an image of the typical landscape/habitat inhabited bymembers of the genus.

Within each species account, information is presented in the following format:

Genus (Subgenus) species name/ author/ dateCommon name: These names are derived for the most part from the Latin

epithet, patronyms, or geographic references. In some cases, I have used the authorof the species name in place of the foregoing. Common names are essential forpublic access to information and particularly in the realm of conservation practices.This is not a new practice; Brullé (1836,1837), for example, provided a commonname in French preceding each of his new species and provided common names forthose already described by his contemporaries and predacessors.

Synonym list: Here I give the original synonymical names arranged in chrono-logical order following the currently accepted name as it was originally proposed. Thename is followed by author and date:page citation.

Distribution: Here, information about Realms, introductions, countries, and insome 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 2200meters; mountains, 2201 to >> n meters. The altitudinal data come directly fromspecimen labels, or from Google look-ups of specimen localities, if data were notfound in publications. Label data given in “feet” were converted to metric.

MICROHABITAT: Under this heading is provided as much detailed information asis known from the literature, field notes, or specimen labels regarding strata (grounddwelling, canopy dwelling, etc.), soil types and general moisture content, near or farfrom water, snow, or other physical feature of the environment wherein members ofthe species are found.

DISPERSAL ABILITIES: Wing length reference; flying, running, burrowing, andclimbing 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 personalcommunications from other Carabidologists.

In the following species accounts, a hyphen (-) is used for its normal purpose; adash (–) is used to indicate a continuum between entries (e.g. between measures,altitudinal ranges, or across months), and for separation between Country and itsStates/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,

CuratorCMNH, Carnegie Museum of Natural History, Pittsburg, PA, Robert Davidson,

CuratorIADIZA, Instituto Argentino de Investigaciones de las Zonas Áridas, Mendoza,

Argentina, Sergio Roig-Juñent, Director and CuratorNMNH, National Museum of Natural History, Smithsonian Institution, Terry L.

Erwin, CuratorUASM, 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 ofany errors, omissions, bibliographic additions, or supply new data and/or images, sothat these may be included both in the volume supplementals and on the websitedatabase. 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 taxonomyof all carabid species described for the Western Hemisphere plus many articles on wayof life. With each ensuing volume, a supplemental bibliography will be added to fill inthe 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 M420microscope with a JVC Digital Camera KY-F7OB attached and processed with GTVision™ software (refered to in the captions as “extended focus image,” or for larger

METHODS | 13


adults, imaged with a Microptics ™ Digital Lab XLT (refered to in the captions as“digital image”.) Leg and antennae orientation differs among habitus images becauseall were reoriented after removal from pins or points and pin holes rendered inPhotoshop™. Background color varies depending on the colors of the specimenbeing imaged. Figure captions of montage and Microptics imaged specimens includean ADP or other number which is an unique label identification number for thespecimen that was illustrated. This number links the specimen and associated illustrationand/or images to additional information in electronic databases at the NMNH. Imagesprovided 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 freshlydead specimen posed for the image. Plate captions include a measure indicating theABL (apparent body length, from Ball, 1982) of the specimen montaged or digitallyimaged with the Microptics setup. Images of live beetles do not have a scale. Thesecaptions also include locality information for individuals imaged, as well as a referencedexemplar landscape/habitat typical for members of the genus. Image credits areprovided in each caption. All extended focus images and Microptics images wereproduced and rendered as necessary in Photoshop by Warren E. Steiner Jr., with theexception of Notiokasis (see caption).

Acknowledgements

First and foremost, heartfelt and gracious thanks go to George E. Ball for myapprenticeship into the world of carabid beetles and his continuing role as my mentorand amigo; and, to his colleague and fellow Carabidologist, Danny Shpeley, whosededicated work on the Méxican collection (UASM) contributed significantly to thisTreatise; and, to Andre L. Larochelle and Marie Claude Larivière for stimulating thiseffort 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. ToGrace 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 intothis 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 ofNewfound, and later in his Magnum Opus of Canada and Alaska in which he presenteda 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 anddedicated it to Carl; I captured their enhancement and built on it. Our debt to Carl isenormous. Special thanks also go to George Venable who got me into using FileMakerProand the resulting data base on which this Treatise is built; to Warren E. Steiner whoprovided the extended focus images and Microptics images for this volume plus manytrips to the library, and to Stephen McJonathan of GT Vision for the loan of equipmentand software necessary to make the extended focus images; to Roy Anderson, AllanAshworth, Christopher J. Earle, Henri Goulet, David Kavanaugh, David Maddison,and Grace Servat who provided other images which are credited appropriately in thefigure captions; to many colleagues who sent their publications to the SmithsonianCarabid Library; and to David H. Kavanaugh, Roberta Brett, and Allen Ashworth fortheir valuable contributions to the Nebriini, Omophronini, and Trachypachidaesections, respectively. And, a hearty thanks to Sergio Roig-Juñent who personallychecked the identifications of Ceroglossus species in the NMNH and to Soledad Sallenavewho did some library work for the Patagonian references.

ACKNOWLEDGEMENTS | 15


Arrangement of Western Hemisphere taxa1

Caraboidea

Volume 1

Family TRACHYPACHIDAE

Tribe Systolosomatini, SystolosomaTribe Trachypachini, Trachypachus

Family CARABIDAE

Division NEBRIIFORMESSubfamily CARABINAESupertribe NEBRIITAETribe Pelophilini, PelophilaTribe Nebriini, Leistus, Nebria, NippononebriaTribe Notiokasiini, NotiokasisTribe Opisthiini, OpisthiusTribe Notiophilini, NotiophilusSupertribe CICINDITAETribe Cicindini, CicindisSupertribe OMOPHRONITAETribe Omophronini, OmophronSupertribe LORICERITAETribe Loricerini, LoriceraSupertribe CARABITAETribe Carabini, Carabus, Calosoma, Callisthenes, Calopachys, Ceroglossus[Tribe Pamborini]Tribe Cychrini, Cychrus, Scaphinotus, SphaeroderusSupertribe CICINDELITAETribe Collyridini, Ctenostoma

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

SPECIES ACCOUNTS / TRACHYPACHIDAE | 17

SPECIES ACCOUNTS

TRACHYPACHIDAEFALSE GROUND BEETLES

Systolosomatini Erwin 1985NEAUSTRAL 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 1849NEAUSTRAL FALSE GROUND BEETLES

(Plate 1)

Systolosoma Solier, 1849:241Notioxenus 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 1849SHORT NEAUSTRAL FALSE GROUND BEETLE

Systolosoma breve Solier, 1849:242Notioxenus 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 Valdivianrainforest with Nothofagus dombeyi (Mirb.) Blume and Araucaria araucana (Molina) K. Koch, Coigue-Maniu Forest, and Nire/Steppe. MICROHABITAT: On sandy alluvium and open sand, as well asin 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-baitedpitfall traps. They fly low to the ground and are often caught in window traps.


References: Ashworth (pers. comm.), Erwin (2001), Data from CAS, NMNH collections

Systolosoma lateritium Nègre 1973TILED 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 rainforests, 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), andat 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 beenfound in January, March, and October – December. BEHAVIOR: Adults take cover in leaf litterand are attracted to dung-baited pitfall traps.References: Ashworth (pers. comm.), Erwin (2001), Nègre (1973), Data from CAS, NMNHcollections

Trachypachini Thomson 1857TEMPORAL 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 1844ROUGH TEMPORAL FALSE GROUND BEETLES

(Plate 2)

Trachypachus Motschulsky, 1844:86Trachypachys 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 facedembankments of exfoliating clay.References: Erwin (2001), Lindroth (1961), Maddison (2001)

SPECIES ACCOUNTS / TRACHYPACHIDAE | 19

Trachypachus gibbsii LeConte 1861GIBBS’ TEMPORAL FALSE GROUND BEETLE

Trachypachus Gibbsii LeConte 1861:339Trachypachus californicus Motschulsky, 1864:194Trachypachus alticola Casey, 1920:144

Distribution: Native, New World. CANADA – BC; USA – CA, ID, MT, OR, WA.Way of Life: MACROHABITAT: Midlands to mountains, 762 – 3505 meters altitude.MICROHABITAT: Adults of this species are stenotopic, restricted to a special habitat where itoften is abundant: on the banks of large rivers and small forested creeks, often some distancefrom the water. Also, they occur under poplars (Populus balsamifera L.) where the ground, halfshaded, consists of fine dusty dry sand covered with dry leaves, no herb layer present, and inpine/fir forests in similar soil conditions. DISPERSAL ABILITIES: Macropterous, and recorded inflight. SEASONAL OCCURRENCE: Adults have been found in March, May – September, andNovember. BEHAVIOR: Adults run in the sunshine on open bare spots; they have been extractedfrom Noble Fir (Abies procera Rehd.) duff using Berlese funnels.References: Erwin (2001), Lindroth (1961), Data from CAS, NMNH collections

Trachypachus holmbergi von Mannerheim 1853HOLMBERG’S TEMPORAL FALSE GROUND BEETLE

Trachypachus Holmbergi von Mannerheim, 1853:119Trachypachus inermis Motschulsky, 1864:194Trachypachus oregonus Casey, 1920:145Trachypachus 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 regularly-tilled 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, occurringindependent of water and are found on open or slightly shaded ground, usually moraine, withsparse 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 1925SLEVIN’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


running on the cliff above the sandy beach of the Pacific Ocean. The beetles run in the talusslope, consisting of small particles, at the base of the cliff of exfoliating clay. The beetles runalong for a short way in the sunshine, then dive into the pile of debris; they also are foundhigher 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

CARABIDAEGROUND BEETLES

NEBRIIFORMES, PART 1CARABINAENEBRIITAE

Pelophilini Kavanaugh 1996MUD-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 1826MUD-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 situationson soft ground with a rich content of organic matter.References: Erwin (2001), Lindroth (1961), Kavanaugh (1996, 1999)

Pelophila borealis (Paykull) 1790BOREAL MUD-LOVING BEETLE

Carabus borealis Paykull, 1790:61Pelophila elongata, von Mannerheim, 1823:38Pelophila Eschscholtzii, von Mannerheim, 1823:40

SPECIES ACCOUNTS / CARABIDAE | 21

Pelophila Gebleri, von Mannerheim, 1823:38Pelophila marginata, von Mannerheim, 1823:38Pelophila Arctica, Dejean, 1826:265Pelophila Dejeanii, Dejean, 1826:265Pelophila laevigata, Motschulsky, 1844:92Pelophila californica, Motschulsky, 1844:93Pelophila angusticollis, Motschulsky, 1844:98Pelophila ochotica, R.F. Sahlberg, 1844:17Pelophila costata, Ménétriés, 1851:49Pelophila Ulkei, G. Horn, 1870:105Pelophila 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 – 590meters 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, occurringalso above tree line and on the true tundra. Found on soft ground with rich content of organicmatter, usually in dense vegetation (e.g. Juncus balticus sitchensis, Carex aquatilis, Eriophorumangustifolium, Arctophila fulva & Hippuris, sometimes in moss carpets of Campylium stellatum, &co). DISPERSAL ABILITIES: Macropterous. Flight recorded in Europe. Adults and larvae are fastrunners; 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; adultstake cover during the day under vegetal debris, stones, and in moss carpets. They are able towithstand freezing temperatures (as low as -5°C). They have been found over wintering incavities 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) 1863WILD 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 withAlnus rugosa (Du Roi) Sprengel, Salices, & co. MICROHABITAT: Adults are found in dense, wetcarpets of Drepanocladus aduncus, with Naumburgia thyrsiflora, Triglochin palustre, Eleocharis palustris,Carices, Juncus, & co. at the margins of eutrophic marshes; marshy borders of slow streams, inswamps, 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


Nebriini Castelnau de Laporte 1834GAZELLE 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 ofgenera recognized. Adelphotaxon: (Notiokasini + Notiophilini + Opisthiini)Geographic status: Holarctic.References: Erwin (2001), Lindroth (1961).Notes: Presentation of this tribe was significantly enhanced by published and unpublishedinformation shared by David H. Kavanaugh, CAS, as well as suggestions for many of thecommon names.

Leistus Frölich 1799BASKET-THROATS

(Plate 4)

Leistus Frölich, 1799:9Leistidius K. Daniel, 1903:171Leistophorus Reitter, 1905:213Euleistulus Reitter, 1905:224Acroleistus Reitter, 1905:225Neoleistus 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 nosynthesis 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 waterin open situations on soft ground with a rich content of organic matter, or in moistgravelly 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) 1758PALEARCTIC RUSTY BASKET-THROAT

Carabus ferrugineus Linné, 1758:415Carabus spinilabris Panzer, 1797: no. 11Leistus testaceus Frölich, 1799:8Leistus kulti Smetana, 1952:234

Distribution: Introduced. PALEARCTIC; CANADA – NF.


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 openwoodlands. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor 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 knownEuropean species.References: Erwin (1970, 2001), Lindroth (1961), Larochelle & Larivière (2003), Data fromUASM collection

Leistus (Neoleistus) ferruginosus von Mannerheim 1843NEARCTIC RUSTY BASKET-THROAT

Leistus ferruginosus von Mannerheim, 1843:187Leistus Ferrugineus Dejean, 1831:569Leistus 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 deciduousforest. MICROHABITAT: Adults occur on moderately moist, half-shaded ground, usually nearrunning water in leaf litter under Alnus and Salix. DISPERSAL ABILITIES: Macropterous, probablycapable of flight. Adults are fast runners and good climbers. SEASONAL OCCURRENCE: Adultsare found in February, April – October; teneral adults in June in BC. BEHAVIOR: Adults climbon 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 1920LONG-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, inconiferous 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 vagilitylimited 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 inMarch, and May – August; teneral adults in June – July. BEHAVIOR: They are likely larvalhibernators, as are all known European species.References: Erwin (1970, 2001), Kavanugh (in litt.), Larochelle & Larivière (2003), Data fromCAS, NMNH collections


Leistus (Neoleistus) madmeridianus Erwin 1970MAD 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, inconiferous 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 litterunder 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 goodclimbers. 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 1802GAZELLE BEETLES

(Plate 5)

Nebria Latreille, 1802:221Boreonebria 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 isa 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 than335 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 seabeaches, 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 1961SHARP-SHOULDERED GAZELLE BEETLE

Nebria acuta acuta Lindroth, 1961:71

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


Way of Life: MACROHABITAT: Lowland to upland forests, subalpine and alpine zones, 10 –2130 meters altitude. MICROHABITAT: Adults are markedly hygrophilous, usually confined tostony, barren margins of running water, or near snow patches and rills. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inMay – 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 suggestedby D.H. Kavanaugh (pers. comm.) because Lindroth was referring to the acute humeri found inthe adults. Kavanaugh will be using this name in his upcoming publications on this genus.

Nebria (Reductonebria) acuta quileute Kavanaugh 1979QUILEUTE 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, probablycapable 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 1981SONORA 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: Streamsbanks 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 1931SOUTHERN 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:


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 1913ARKANSAS 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 abovetreeline in Canada, 2010 – 3810 meters altitude. MICROHABITAT: Confined to stony-gravelly marginsof fast running waters, usually small brooks; DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJune – 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 1979EDWARDS’ 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: MACROHABITAT: 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 coverin 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 1924FRAGILE GAZELLE BEETLE

Nebria arkansana fragilis Casey, 1924:21Nebria uinta Kavanaugh, 1979:102Nebria 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 nocturnaland carnivorous, adults take cover in the day under stones and drift.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)


Nebria (Reductonebria) arkansana oowah Kavanaugh 1979LA 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: Confinedto margins of cool streams on wet ground. DISPERSAL ABILITIES: Macropterous, probably capableof flight. SEASONAL OCCURRENCE: Adults have been found in August. BEHAVIOR: Adults arenocturnal, carnivorous.References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003),Lindroth (1961)

Nebria (Nebria) brevicollis (Fabricius) 1792SHORT-COLLARED GAZELLE BEETLE

Carabus brevicollis Fabricius, 1792:150Carabus cursor O.F. Müller, 1776:78Carabus rufipes Goeze, 1777:622Carabus infida Rossi, 1792:88Nebria rugimarginata Marsham, 1802:444Nebria fuscata Bonelli, 1810:64Nebria lata Newman, 1833:284Nebria varicornis Newman, 1833:285Nebria elongata Fischer von Waldheim, 1828:266Nebria rufomarinata Fischer von Waldheim, 1828:248Nebria Tamsii Dejean, 1831:573Nebria barbara Chaudoir, 1843:748Nebria rufipes Dalla Torre, 1877:24Nebria cursor Bedel, 1879:17Nebria sicula Preudhomme de Borre & Ragusa, 1882:181Nebria obscurata J.R. Sahlberg, 1903:3Nebria 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 orwoodland habitats such as deciduous forests and parks and often invades adjacent agriculturallands. They prefer moist mull ground in beech forests. It is one of the most abundant speciesin 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 moraineof 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 ashort 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)


Nebria (Catonebria) calva Kavanaugh 1984MOUNT 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 coldstreams in upper forested and treeline areas. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJuly. BEHAVIOR: Adults are nocturnal, carnivorous.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nebria (Catonebria) carri Kavanaugh 1979CARR’S GAZELLE BEETLE

Nebria carri Kavanaugh, 1979:107

Distribution: Native, New World. USA – ID, MT.Way of Life: MACROHABITAT: 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: Adultshave been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nebria (Catonebria) catenata Casey 1913SOUTHERN CHAIN-BACKED GAZELLE BEETLE

Nebria catenata Casey, 1913:49

Distribution: Native, New World. USA – CO, UT.Way of Life: MACROHABITAT: 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 1961GRAHAM 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 thatare at least partially shaded by the adjacent forest edge. MICROHABITAT: Stenotopic adults haveonly been found where the cobble is piled at least six inches deep and freshwater seeps through


the sand/gravel substrate below. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagilitylimited 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 adultsare 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 1943COLORADO GAZELLE BEETLE

Nebria coloradensis Van Dyke, 1943:19

Distribution: Native, New World. USA – CO.Way of Life: MACROHABITAT: Mountains, 2870 – 3350 meters altitude. MICROHABITAT: Foundon wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 1925THICK-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 nearsnow patches on moist or wet ground. DISPERSAL ABILITIES: Macropterous, probably capableof 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 1949INTERMEDIATE GAZELLE BEETLE

Nebria crassicornis intermedia Van Dyke, 1949:49

Distribution: Native, New World. CANADA – AB, BC; USA – ID, MT, OR, UT, WA, WYWay 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 inopen 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 offlight. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Adultsare nocturnal, carnivorous. Hibernation takes place in both the adult and larval stages.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003),Lindroth (1961)


Nebria (Reductonebria) danmanni Kavanaugh 1981DAN 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: Confinedto 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 ofthe glaciers preying on half frozen insects. Adults are nocturnal, carnivorous, taking cover inthe day under glaciers in rubble.References: Erwin (2001), Kavanaugh (1981, 1985, in litt.), Larochelle & Larivière (2003)

Nebria (Reductonebria) darlingtoni Kavanaugh 1979DARLINGTON’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 riverbanks and near waterfalls on wet ground. DISPERSAL ABILITIES: Macropterous, probably capableof 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 1971DESOLATE GAZELLE BEETLE

Nebria desolata Kavanaugh, 1971:41

Distribution: Native, New World. USA – UT.Way of Life: MACROHABITAT: Upland desert, 1710 – 2130 meters altitude. MICROHABITAT: Atthe 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 arenocturnal, 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 1863SEA BEACH GAZELLE BEETLE

Nebria diversa LeConte, 1863:2Nebria livida LeConte, 1859:84Nebria Townsendi Casey, 1924:19

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


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 larvaeare found beneath driftwood and other debris above the highest tide line, especially where sanddunes have developed above the beaches. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary – 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 thewater. Larvae hide under driftwood. Larvae hibernate.References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003),Lindroth (1961)

Nebria (Reductonebria) eschscholtzii Ménétriés 1843ESCHSCHOLTZ’S GAZELLE BEETLE

Nebria Eschscholtzii Ménétriés, 1843:55Nebria tenuipes Casey, 1913:51Nebria transversa Casey, 1920:152Nebria formalis Casey, 1920:153Nebria pallidissima Casey, 1924:19Nebria pugetana Casey, 1924:19

Distribution: Native, New World. USA – CA, ID, NV, OR, WAWay 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 beenfound in January – December. BEHAVIOR: Adults are nocturnal, carnivorous, and take coverunder 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 1844FRIGID GAZELLE BEETLE

Nebria frigida R.F. Sahlberg, 1844:11Nebria viridis G. Horn, 1870:101Nebria parvula J.R. Sahlberg, 1887:47Nebria 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 adultsalways 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 arenocturnal, carnivorous, taking cover in the day under stones. They occur up to and slightlyabove 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)


Nebria (Catonebria) gebleri albimontis Kavanaugh 1984WHITE 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 onwet 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 1979CASCADE 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: Onriver 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 1979STRAWBERRY 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 offlight. SEASONAL OCCURRENCE: Adults have been found in May and July. BEHAVIOR: Adults arenocturnal, carnivorous, taking cover in the day.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nebria (Catonebria) gebleri gebleri Dejean 1831GEBLER’S GAZELLE BEETLE

Nebria Gebleri Dejean, 1831:573Nebria 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 inMay – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day understones or drift. Adults hibernate.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)


Nebria (Catonebria) gebleri rathvoni LeConte 1853RATHVON’S GAZELLE BEETLE

Nebria Rathvoni LeConte, 1853:400

Distribution: Native, New World. USA – CA, NV.Way of Life: MACROHABITAT: Midlands to mountains, 910 – 3200 meters altitude.MICROHABITAT: On river banks on wet ground. DISPERSAL ABILITIES: Macropterous, probablycapable 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 1979SISKIYOU 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 recordedfrom above tree line. MICROHABITAT: Along mountain streams on forested slopes, often quitesmall brooks, with stony, barren banks; always close to the water. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inJune – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day understones or drift. Larvae hibernate.References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003)

Nebria (Boreonebria) gouleti Kavanaugh 1979GOULET’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 ratherwet 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 1820GREGARIOUS GAZELLE BEETLE

Nebria gregaria Fischer von Waldheim, 1820: pl. 6Nebria cuneata Casey, 1913:50

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


sea beaches, but usually found on open, moderately wet ground on peaty or gravelly groundwith not too dense vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagilitylimited 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) 1837CHESTNUT-LEGGED GAZELLE BEETLE

Helobia castanipes Kirby, 1837:20Nebria moesta LeConte, 1850:209Nebria elias Motschulsky, 1865:276Nebria labradorica Casey, 1920:151Nebria prominens Casey, 1920:151Nebria 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 atthe 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 mixedwith sand or detritus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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 treesand 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 1979LASSEN GAZELLE BEETLE

Nebria gyllenhali lassenensis Kavanaugh, 1979:96

Distribution: Native, New World. USA – CA, OR.Way of Life: MACROHABITAT: 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: Adultshave 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)


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

Nebria gyllenhali lindrothi Kavanaugh, 1979:97

Distribution: Native, New World. USA – AZ, CO, NM, UT, WY.Way of Life: MACROHABITAT: 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 amongstones. Adults hibernate.References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003)

Nebria (Reductonebria) haida Kavanaugh 1984HAIDA 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 toKavanaugh (1992), this species is restricted to the alpine zone. MICROHABITAT: Adults andlarvae are found under stones on ridges and summits above tree line, especially in areas ofsparse vegetation. Unlike most alpine Nebria, members of this species avoid waterside areasand edges of snowfields and are instead most abundant in the driest sites available, moistground with little vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagilitylimited 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 1863HUDSONIAN 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 – 3200meters altitude. MICROHABITAT: On barren, usually stony banks of small and large runningwater, not confined to really cold water. DISPERSAL ABILITIES: Macropterous, probably capableof flight. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Adultsare 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)


Nebria (Catonebria) ingens ingens G. Horn 1870CUP-FOOTED GAZELLE BEETLE

Nebria ingens ingens G. Horn, 1870:98Nebria raveni Van Dyke, 1953:102

Distribution: Native, New World. USA – CA.Way of Life: MACROHABITAT: Mountains, 3350 – 4150 meters altitude. MICROHABITAT: Granitetalus 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 adaptationof the tarsomeres, highly unusal in nebrines.

Nebria (Catonebria) ingens riversi Van Dyke 1925RIVERS’ 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: Marginsof glaciers, snow patches, and glacial streams. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in July – August. BEHAVIOR: Adults are nocturnal, carnivorous.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nebria (Reductonebria) jeffreyi Kavanaugh 1984JEFFREY’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, probablycapable 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 1979BALL’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


flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in May – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in theday under drift.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nebria (Catonebria) kincaidi kincaidi Schwarz 1900KINCAID’S GAZELLE BEETLE

Nebria Kincaidi Kincaidi Schwarz, 1900:525Nebria 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: Adultshave been found in May – September. BEHAVIOR: Adults are nocturnal, carnivorous, takingcover in the day. Adults hibernate.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)

Nebria (Catonebria) labontei Kavanaugh 1984LABONTE’S GAZELLE BEETLE

Nebria labontei Kavanaugh, 1984:163

Distribution: Native, New World. USA – OR.Way of Life: MACROHABITAT: Mountains, 2040 – 2440 meters altitude. MICROHABITAT: Wetground. 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 1979BELLS’ 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 beenfound in April – October. BEHAVIOR: Adults are nocturnal, carnivorous.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)


Nebria (Boreonebria) lacustris lacustris Casey 1913LACUSTRINE 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 mouthof Rock Run on sand mixed with clay, under stones, at the southeast corner of Island on coarsesand mixed with silt, by splashing. DISPERSAL ABILITIES: Macropterous, probably capable offlight. SEASONAL OCCURRENCE: Adults have been found in March – November; teneral adultsin April – May. BEHAVIOR: These beetles hide during the day and forage at night on dead anddying insects along the stream margin. Adults are nocturnal, carnivorous, taking cover in theday 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 1979LITUYA’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 thusvagility limited to walking or running. Climber on fallen trees. SEASONAL OCCURRENCE: Adultshave 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 1984SOUTH 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 beachesthat 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 thesand/gravel substrate below. Some beaches on which this species occurs are strewn withdriftwood. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Adults have been found in June – August; copulatingpairs 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 theday under the cobble. Hibernation, at least in part, as larvae.References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003)


Nebria (Reductonebria) lyelli Van Dyke 1925MOUNT 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 themargins of glacial streams on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJune – 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 1828MANNERHEIM’S GAZELLE BEETLE

Nebria Mannerheimii Fischer von Waldheim, 1828:253Nebria oregona Casey, 1913:52Nebria corvallis Casey, 1924:20Nebria 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 arewidespread 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. innaturally disturbed areas or synanthropic sites such as quarries), and under stones on open,exposed inorganic shores of large lowland streams. DISPERSAL ABILITIES: Macropterous, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in March – November;teneral adults in May – June. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in theday 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 1981GIULIANI’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 theborder of brooks on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable offlight. SEASONAL OCCURRENCE: Adults have been found in June – July. BEHAVIOR: Adults arenocturnal, carnivorous, taking cover in the day.References: Erwin (2001), Kavanaugh (1981, 1985, in litt.), Larochelle & Larivière (2003)


Nebria (Catonebria) meanyi lamarckensis Kavanaugh 1979LAMARCK’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 capableof flight. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Adultsare nocturnal, carnivorous, taking cover in the day.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nebria (Catonebria) meanyi meanyi Van Dyke 1925MEANY’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 inJune – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day understones. Adults hibernate.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)

Nebria (Catonebria) meanyi sylvatica Kavanaugh 1979SYLVAN 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: Nearsprings 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 1820METALLIC GAZELLE BEETLE

Nebria metallica Fischer von Waldheim, 1820: pl. 6Nebria 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 runningwater where they occur some distance from the bank under cover of shrubs and trees, sometimeson clayish ground. On the Alaskan coast and adjacent islands, adults frequent open country


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

Nebria (Reductonebria) navajo Kavanaugh 1979NAVAJO 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 thusvagility 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 1979GASPÉ 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 bareground that is wet and stony. DISPERSAL ABILITIES: Macropterous, probably capable of flight.SEASONAL OCCURRENCE: Adults have been found in June – September; copulating pairs in lateJuly; teneral adults in June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking coverin 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) 1790SNOW GAZELLE BEETLE

Carabus nivalis Paykull, 1790:52Nebria bifaria von Mannerheim, 1853:128Nebria femoralis Motschulsky, 1859:541Nebria mollis Motschulsky, 1866:274Nebria molbis Motschulsky, 1866:274Nebria femorata Motschulsky, 1866:275Nebria obscuripes Poppius, 1906:19Nebria 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


margins of snowfields north of, and above tree line, but adults follow rivers and big brookswith cold water, in particular glacier outflows, deep down into the forest region. They alwayskeep 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 beenfound 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 1979CHUSKA 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in July – August. BEHAVIOR: Adults are nocturnal, carnivorous. Adults take cover understones.References: Erwin (2001), Kavanaugh (1979, 1985, in litt.)

Nebria (Reductonebria) obliqua obliqua LeConte 1866OBLIQUE GAZELLE BEETLE

Nebria obliqua LeConte, 1866:363Nebria obtusa LeConte, 1878:478Nebria incerta Casey, 1913:53Nebria testaceipes Casey, 1913:54Nebria 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; theyare not dependent on cold, clear water and are mostly found on silty places where the river runsslowly. 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 1878OVAL GAZELLE BEETLE

Nebria ovipennis LeConte, 1878:477

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


Way of Life: MACROHABITAT: Uplands to mountains, and subalpine zone, 2040 – 3510 metersaltitude. 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 placein both larval and adult forms.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)

Nebria (Reductonebria) pallipes Say 1823PALE-LEGGED GAZELLE BEETLE

Nebria pallipes Say, 1823:78Nebria 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. lacustrisunder 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 inAugust; 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 likelyboth 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 1931PARADISE GAZELLE BEETLE

Nebria paradisi Darlington, 1931:104Nebria kincaidi Bänninger, 1925:264Nebria columbiana Van Dyke, 1926:10Nebria vandykei Darlington, 1930:24

Distribution: Native, New World. USA – OR, WA.Way of Life: MACROHABITAT: Midlands to mountains, alpine and subapline zones, 1160 – 2440meters altitude. MICROHABITAT: Moderately hygrophilus and found on margins of streams andsnowfields on rather wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility 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)


Nebria (Catonebria) piperi Van Dyke 1925PIPER’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 metersaltitude. MICROHABITAT: Under large stones near the water on barren, stony banks of riverswith cold water. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL

OCCURRENCE: Adults have been found in May – October; tenerals in June. BEHAVIOR: Adultsare 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 1972PIUTE 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 1984SEVIER 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: Wetground 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 1979UTAH GAZELLE BEETLE

Nebria trifaria utahensis Kavanaugh, 1979:110

Distribution: Native, New World. USA – UT.Way of Life: MACROHABITAT: Mountains, 2290 – 2500 meters altitude. MICROHABITAT: Wetground 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 (1979, 1985, in litt.)

Nebria (Catonebria) purpurata LeConte 1878PURPLE GAZELLE BEETLE

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

Distribution: Native, New World. USA – CO, NM.Way of Life: MACROHABITAT: Mountains, subalpine and alpine zones, 2350 – 3810 metersaltitude. MICROHABITAT: Along cold mountain streams on wet sandy ground. DISPERSAL


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 goodswimmers 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 1979MODOC GAZELLE BEETLE

Nebria sahlbergii modoc Kavanaugh, 1979:99

Distribution: Native, New World. USA – CA.Way of Life: MACROHABITAT: Uplands, 1740 – 1930 meters altitude. MICROHABITAT: Alongbrooks 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 1828SAHLBERG’S GAZELLE BEETLE

Nebria Sahlbergii Fischer von Waldheim, 1828:254Nebria violacea Motschulsky, 1850:73Nebria 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 onshady wet gravelly-stony ground. DISPERSAL ABILITIES: Macropterous, probably capable offlight. SEASONAL OCCURRENCE: Adults have been found in May – August; teneral adults inMarch – November. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the dayunder stones. Likely both adults and larvae hibernate.References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003)


Nebria (Reductonebria) sahlbergii triad Kavanaugh 1979TRINITY 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 capableof 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 1979BEVERLY 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 bankson 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 1925SCHWARZ’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: Alongstream margins on wet bare coarse sand. DISPERSAL ABILITIES: Macropterous, probably capableof flight. SEASONAL OCCURRENCE: Adults have been found in June – October. BEHAVIOR: Adultsare 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 1984SIERRA BLANCA GAZELLE BEETLE

Nebria sierrablancae Kavanaugh, 1984:164

Distribution: Native, New World. USA – NM.Way of Life: MACROHABITAT: Mountains, 2740 – 3510 meters altitude. MICROHABITAT: Shadedwet 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.)


Nebria (Catonebria) spatulata sierrae Kavanaugh 1979SIERRAN 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 1925SPATULATE GAZELLE BEETLE

Nebria spatulata Van Dyke, 1925:119Nebria fusiformis Van Dyke, 1926:11

Distribution: Native, New World. USA – CA.Way of Life: MACROHABITAT: Mountains, alpine zone, 3050 – 3140 meters altitudeMICROHABITAT: 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 1984STEENS MOUNTAINS GAZELLE BEETLE

Nebria steensensis Kavanaugh, 1984:165

Distribution: Native, New World. USA – OR.Way of Life: MACROHABITAT: Uplands to mountains, 2190 – 2725 meters altitude.MICROHABITAT: Margins of small streams. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJuly – September. BEHAVIOR: Adults are nocturnal, taking cover in the day; they are carnivorous.References: Erwin (2001), Kavanaugh (1985, in litt.)

Nebria (Reductonebria) suturalis LeConte 1850SEAMED GAZELLE BEETLE

Nebria suturalis LeConte, 1850:209Nebria longula LeConte, 1878:478Nebria nimbosa Casey, 1920:150

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


Way of Life: MACROHABITAT: Midlands to mountains, subalpine, and alpine zones, 1190 –4350 meters altitude. MICROHABITAT: Confined to mountain tops; adults are found under stoneswhere water is seeping or trickling down the mountain side in open places above tree line. Someindividuals 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 1984PASQUINEL’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: Nearstreams and lakes. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 1878NORTHERN CHAIN-BACKED GAZELLE BEETLE

Nebria trifaria LeConte, 1878:478Nebria tetonensis Erwin & Ball, 1972:95

Distribution: Native, New World. USA – ID, MT, NV, UT, WY.Way of Life: MACROHABITAT: 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 arenocturnal, carnivorous, taking cover in the day.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)

Nebria (Boreonebria) turmaduodecima Kavanaugh 1981TROUP 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 flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inAugust. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in day under stones.References: Erwin (2001), Kavanaugh (1981, 1985, in litt.), Larochelle & Larivière (2003)


Nebria (Catonebria) vandykei vandykei Bänninger 1928VAN 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 – 2040meters 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 1979WYEAST’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


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 1984WALLOWA 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, probablycapable 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 1979OASIS 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 brookmargins. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Adults arenocturnal, carnivorous, taking cover in the day.


References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nebria (Reductonebria) zioni zioni Van Dyke 1943ZION 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 metersaltitude. MICROHABITAT: On brook margins. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inMay – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day.References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

Nippononebria Uéno 1955FALSE GAZELLE BEETLES

(Plate 6)

Nippononebria Uéno, 1955:49Vancouveria 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 1799Type 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) 1984HIGH 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)


Nippononebria (Vancouveria) campbelli (Kavanaugh) 1984CAMPBELL’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: Accordingto Kavanugh (in litt.), well above treeline at Mount Baker, under a stone on a rocky outcropsurrounded by huge snowfields on one of the mountain’s main ridges. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inJuly – 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) 1870GREENISH FALSE GAZELLE BEETLE

Nebria virescens G. Horn, 1870:10Nebria 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: Adultshave been found in February – December. BEHAVIOR: Adults are nocturnal, carnivorous, takingcover in the day. Adults, at least in part, hibernate.References: Erwin (2001), Kavanaugh (in litt.), Larochelle & Larivière (2003), Lindroth (1961)

Notiokasiini Kavanaugh & Nègre 1983SOUTHERN SISTER BEETLES

Notiokasiini Kavanaugh & Nègre, 1983:551

Number of worldwide genera: One.Number of Western Hemisphere genera: One.Taxonomy: Stable. Adelphotaxon: Notiophilus Dumeril 1806 + OpisthiniGeographic status: NeotropicalDistribution: Native, New World. Brazil; Uruguay.References: Erwin (2001), Kavanaugh & Nègre (1983)

Notiokasis Kavanaugh & Nègre 1983SOUTHERN 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.


Taxonomy: Stable. Adelphotaxon: See Notiokasiini, above.Type species: Notiokasis chaudoiri, Kavanaugh & Nègre, 1983Distribution: 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 1983CHAUDOIR’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: Adultshave been found in July. BEHAVIOR: Unknown.References: Erwin (2001), Kavanaugh & Nègre (1983)

Opisthiini Dupuis 1912FALSE MARSH AND BOG BEETLES

Opisthiini Dupuis 1912:1

Number of worldwide genera: Two.Number of Western Hemisphere genera: One.Taxonomy: Stable. Adelphotaxon: Notiokasini + NotiophiliniGeographic status: Holarctic.References: Erwin (2001), Lindroth (1961), Kavanaugh (1996, 1999)

Opisthius Kirby 1837FALSE 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 1920Type species: Opisthius richardsoni Kirby, 1837Distribution: Native, New World (one species).Habitat: Banks of rivers and big brooks.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961)


Opisthius richardsoni Kirby 1837RICHARDSON’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, consistingof clay, often mixed with sand and gravel. DISPERSAL ABILITIES: Macropterous, capable offlight, 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 theStaphylinds of the genus Bledius. Adults keep close to the water where the ground is free ofvegetation. 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 1850BIG-EYED BEETLES

Notiophilini Mostschulsky, 1850:16

Number of worldwide genera: One.Number of Western Hemisphere genera: One.Taxonomy: Stable. Adelphotaxon: Opisthini Dupuis 1912Geographic status: Holarctic, Neotropical.References: Erwin (2001), Barsevskis (1994), Lindroth (1961)

Notiophilus Dumeril 1806BIG-EYED BEETLES

(Plate 9)

Notiophilus Dumeril, 1806:193Nothiophilus Dumeril, 1806:194Nothiophilus Say, 1823:81Latviaphilus Barsevskis, 1994:1Makarovius 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 ratherxerophilus. Most species occur in open country where the vegetation is sparse and the groundconsists of gravel, others belong to open forested areas. Adults are pronouncedly carnivorous,


seen feeding on gants and spiders. Wing dimorphism occurs more regularly in this genus thanany other large carabid genus.References: Erwin (2001), Barsevskis (1994), Lindroth (1961)

Notiophilus aeneus (Herbst) 1806BRASSY BIG-EYED BEETLE

Elaphrus aeneus Herbst, 1806:235Notiophilus 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 androadsides. MICROHABITAT: At Plummers Island, Maryland, adults were found in moss and deepleaf 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 oftrees. SEASONAL OCCURRENCE: Adults have been found in January – December. BEHAVIOR:Adults are mostly diurnal and active in the sunshine; they are carnivorous, probably predaceouson 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) 1758LINNÉ’S BIG-EYED BEETLE

Cicindela aquaticus Linné, 1758:408Cicindela pusillus Schreber, 1759:10Elaphrus semipunctatus Fabricius, 1775:227Notiophilus metallicus Waterhouse, 1833:203Notiophilus parvulus Waterhouse, 1833:208Notiophilus dauricus Motschulsky, 1860:539Notiophilus strigifrons Baudi di Selve, 1864:146Notiophilus Hardyi Putzeys, 1866:165Notiophilus apicalis Dalla Torre, 1877:22Notiophilus obscurus Dalla Torre, 1877:22Notiophilus obsoletus Dalla Torre, 1877:22Notiophilus Dybowskii Lomnicki, 1903:113Notiophilus Kroli Lomnicki, 1903:113Notiophilus pristinus Friederichs, 1903:88Notiophilus Semenovi Tschitschérine, 1903:116Notiophilus Blacki Edwards, 1913:70Notiophilus Spaethi Munster, 1922:140Notiophilus aquaticus bimaculatus Barševskis, 1993:51Notiophilus aquaticus pseudoaesthuans Barševskis, 1994:7


Notiophilus aquaticus stoschkae Barševskis, 1994:7Notiophilus 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 coastaltundra, sea level – 3048 meters altitude. MICROHABITAT: On moraines, fields, meadows, vacantlots, cultivated fields (e.g. corn, barley), roadsides, sand pits, gravel pits, forest edges, and inmixed deciduous forests. Adults occur on open ground that is well-drained, and consisting ofmoderately 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 prescribed-burned forest. DISPERSAL ABILITIES: Wing-dimorphic. European records indicate macropterousadults fly. SEASONAL OCCURRENCE: Adults have been found in January – December; teneraladults 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) 1779SPOTTED BIG-EYED BEETLE

Elaphrus biguttatus Fabricius, 1779:232Dermestes aquaticus Geoffroy, 1785:21Elaphrus semipunctatus Duftschmid, 1812:90Notiophilus latus Waterhouse, 1833:209Notiophilus nitidus Waterhouse, 1833:210Notiophilus striatus Waterhouse, 1833:209Notiophilus lateralis Motschulsky, 1864:192Notiophilus melanophthalmus Slosser-Klekovski, 1877:12Notiophilus pseudoquadripunctatus Everts, 1918:43Notiophilus pseudolateralis Louvet, 1925:6Notiophilus 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 occursin 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 drygravelly, 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 coniferneedles or pieces of wood; they are carnivorous and in Europe, adults have been observed eatingCollembola, mites, flies, and other arthropods. Larvae are known to climb on tree trunks. Thisspecies 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


Notiophilus borealis Harris 1869NORTHERN 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 – 2332meters altitude. MICROHABITAT: On open or slightly shaded rather dry, gravelly or sandy groundcovered 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 beenfound in June – October; tenerals in August – September. BEHAVIOR: Mostly diurnal, active in thesunshine; 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 1913CHIHUAHUA 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/pineforests and at higher altitudes in oak/pine/fir forests. MICROHABITAT: In leaf litter, particularlyconifer litter. DISPERSAL ABILITIES: Brachypterous (at least in part), hence flightless thus vagilitylimited 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 1920STRAIGHT BIG-EYED BEETLE

Notiophilus directus Casey, 1920:142Notiophilus 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 byshrubs or trees, or, covered with tiny mosses on moraines; or in the vicinity of brooks andlakes. DISPERSAL ABILITIES: Wing-dimorphic: macropterous or brachypterous. Macropterousindividuals 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


Notiophilus intermedius Lindroth 1954INTERMEDIATE 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 onriver banks. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor 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 daysunder mosses, lichens, or low plants, e.g. Empetrum (Ericaceae).References: Erwin (2001), Larochelle & Larivière (2003)

Notiophilus nemoralis Fall 1906WOODLAND 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 oncloudy 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 1857BRIGHT 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 opendry, 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 inMay – June. BEHAVIOR: Mostly diurnal, xerophilus, active in the sunshine; adults take cover oncloudy 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 1848NINE-LINED BIG-EYED BEETLE

Notiophilus novemstriatus LeConte, 1848:350Notiophilus cribrilaterus Motschulsky, 1864:193


Notiophilus quadrifoveolatus T.W. Harris, 1869:212Notiophilus 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 dryground, 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: Wing-dimorphic: brachypterous or macropterous. According to Lindroth (1961) macropterousindividuals 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 ofgrass. 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) 1812MARSH BIG-EYED BEETLE

Elaphrus palustris Duftschmid, 1812:192Notiophilus nitidulus Waterhouse, 1833:203Notiophilus parallelus Waterhouse, 1833:204Notiophilus newmanni Waterhouse, 1833:205Notiophilus Davisi Waterhouse, 1833:205Notiophilus brevis Waterhouse, 1833:206Notiophilus tibialis Waterhouse, 1833:206Notiophilus latior Waterhouse, 1833:207Notiophilus melancholicus Dalla Torre, 1877:23Notiophilus hirticollis Chaudoir, 1882: LXXXNotiophilus atavus Friedrichs, 1903:88Notiophilus decemstriatus Roubal, 1948:45Notiophilus impunctatus Pulpan, 1948:49Notiophilus 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 cultivatedfields 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 driftmaterial. SEASONAL OCCURRENCE: Adults have been found in Canada in June – July; teneraladults 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


Notiophilus rufipes Curtis 1829RUFUS-LEGGED BIG-EYED BEETLE

Notiophilus rufipes Curtis, 1829:254Notiophilus fulvipes Motschulsky, 1845:12Notiophilus rufipes Chaudoir, 1844:439Notiophilus 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 groundnear the base of separately growing trees. DISPERSAL ABILITIES: Wing-dimorphic, mostlymacropterous, capable of flight. SEASONAL OCCURRENCE: In Europe, March, July – August,October. BEHAVIOR: Adults have been observed hibernating in Europe. Adults move fast byrunning and flying, hence most are macropterus.References: Erwin (2001), Barševskis (2004), Larochelle & Larivière (2003), Data from NMNHcollectionNotes: The single know specimen from Georgia was collected in August of 1975 at 44 metersaltitude in the Okefenokee Swamp. Whether the species has become established is unknown.

Notiophilus semiopacus Eschscholtz 1833SEMIOPAQUE 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 andseeps, and in foothill canyons along streams, in parks, and on the sea beach in southern CA; inMéxico 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 antnests. Macropterous adults occasionally found at lights at night.References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections

Notiophilus semistriatus Say 1823SEMI-STRIATE BIG-EYED BEETLE

Notiophilus semistriatus Say, 1823:81Notiophilus confusus LeConte, 1848:349Notiophilus punctatus LeConte, 1850:210Notiophilus americanus T.W. Harris, 1869:213Notiophilus 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.


Way of Life: MACROHABITAT: Lowlands to mountains, subalpine and alpine zones, 200 – 3200meters latitude. MICROHABITAT: On open or half-shaded rather dry, gravelly, sandy, rocky or peatyground; covered with sparse low thin vegetation or patches of mosses and lichens on moraines, inabandoned 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 underreindeer 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, UASMcollections

Notiophilus sierranus Casey 1920MOUNTAIN BIG-EYED BEETLE

Notiophilus sierranus Casey, 1920:140Notiophilus obscurus Fall, 1901:207Notiophilus 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 1906FALL’S BIG-EYED BEETLE

Notiophilus simulator Fall, 1906:86Notiophilus 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: MACROHABITAT: Uplands to mountains, 1520 – 3300 meters altitude.MICROHABITAT: On open or slightly shaded sandy bare ground that is nearly bare except forpatches 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 1881MIRROR-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.


Way of Life: MACROHABITAT: Uplands to mountains, 1540 – 3100 meters altitude, inmesophylic and cloud forests, oak, pine/oak forests, pine/alder/manzanita forest edge, andin 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 adultsin April in CH. BEHAVIOR: Running on leaf litter at night in HD. Adults take cover understones and in coniferous litter.References: Erwin (1991, 2001), Data from NMNH, UASM collections

Notiophilus sylvaticus Eschscholtz 1833FOREST BIG-EYED BEETLE

Notiophilus sylvaticus Eschscholtz 1833:24

Distribution: Native, New World. CANADA – BC, QC; USA – AK, CA, ID, NM, OR, WAWay 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 onslightly shaded, gravelly ground, but not in dense forests. Also, occasionally found inGreenhouses. According to Kavanaugh (1992), this species is widespread on the QueenCharlotte Islands in forested areas at all elevations, from near sea level to above tree line onthe 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 theforest floor throughout the archipelago. Above tree line, beetles occur in the driest sites withvery low, scattered vegetation, and they are especially active in sunlight. DISPERSAL ABILITIES:Wing-dimorphic: brachypterous or macropterous, both forms occurring in the samepopulation. 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 daysunder the loose bark of fallen trees, under fallen trees and stones, in moss, and amongconifer needles. Adults are gregarious.References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections

CICINDITAECicindini

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)


Cicindis Bruch 1908FAIRY 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 1908HORN’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 flatsnear vernal pools. MICROHABITAT: Adults occur on wet to inundated clay mixed with salt wherepolygons form when the clay dries out and with no vegetation cover. DISPERSAL ABILITIES: Adultsare fully winged and fly readily at night; adults also are swimmers and divers in vernal pools with17% salinity. SEASONAL OCCURRENCE: Adults are found in December – March; copulating pairsobserved in March; teneral adults in February. BEHAVIOR: This heretofore very rare species wasrecently found to be markedly unusual among carabid beetles because adults surface-swim anddive in search of their prey, an anostracan fairy shrimp, possibly a species of ThamnocephalusCohen. Adult beetles do not run along the shore at the margin of the ponds that are formed in thegreat salt flats like most carabid in this type of habitat elsewhere. However, they do rest in smallburrows during the day away from the water’s edge, and emerge before nightfall (crepuscularbehavior). They then spend the night in search of food or mates swimming on the water surfaceof markedly alkaline water bodies (17% salinity). Tibial natatorial setae, tarsal setal fringes, ventralvestiture, and genal/pronotal planing surfaces plus decumbent and very sharply pointed mandiblesare 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 ofvernal 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 mudadjacent to the pool, or at UV and Mercury Vapor lights; and they sometimes can be obtained inpitfall traps placed very near the water’s edge. Immature stages and hibernation site unknown.References: Erwin (2001), Kavanaugh & Erwin (1991), Erwin & Aschero (2004), Data fromNMNH collection


OMOPHRONITAEOmophronini 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 1802ROUND SAND BEETLES

(Plate 11)

Omophron Latreille, 1802:89Scolytus Fabricius, 1790:221Epactius Schneider, 1791:23Lithophilus Schneider, 1791:23Homophron Fischer von Walheim, 1828:256Epacter Semenov, 1922:38Andromophron Semenov, 1922:40Oligomophron Semenov, 1922:40Paromophron Semenov, 1922:40Homophron Semenov, 1922:41Stenomophron Semenov, 1922:42Istor Semenov 1922:43Prosecon Semenov, 1922:44Phromoon 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 foundin both Nearctic and Neotropical Realms.Habitat: According to Lindroth (1961), all species of this genus are markedly hygrophilousand 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 theday. 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)


Omophron americanum Dejean 1831AMERICAN ROUND SAND BEETLE

Omophron Americanus Dejean, 1831:583Omophron Sayi Kirby, 1837:65Omophron lacustrae Casey, 1897:301Omophron texanus Casey, 1897:302Omophron admiscens Casey, 1913:8Omophron fontinale Casey, 1913:41Omophron iridescens Casey, 1913:41Omophron Lengi Casey, 1920:135Omophron illustre Casey, 1920:136Homophron 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: MACROHABITAT: Lowlands to uplands, 260 – 2195 meters altitude.MICROHABITAT: On open bare, or almost so, wet ground consisting of sand, or sand mixedwith 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 inApril – 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 banksand 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 andthey 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 1912BÄNNINGER’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: Streammargins. 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 1852TOOTHED ROUND SAND BEETLE

Omophron dentatus LeConte, 1852:200

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


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: Adultshave 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). Adultsoccasionally found at lights at night.References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections

Omophron dominicense Chaudoir 1868INSULAR 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 1852GILA RIVER ROUND SAND BEETLE

Omophron gilae LeConte, 1852:201Omophron pallidus Casey, 1897:305Omophron 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 1868PLEASING 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, UASMcollections


Omophron grossum Casey 1909LARGE 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 openwet, 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) 1801LARGE-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 sometimesactive 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 sometimesas 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 1912MEXICAN 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 1848SHINY ROUND SAND BEETLE

Omophron nitidus LeConte, 1848:347Omophron 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.


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 havebeen found in March – December. BEHAVIOR: Nocturnal, adults take cover during the day inburrows 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 1870EFFACED ROUND SAND BEETLE

Omophron obliteratus G. Horn, 1870:73Omophron sonorae Casey, 1897:304Omophron utense Casey, 1913:43Omophron 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 pineforest along creeks in canyons. MICROHABITAT: Open ground on creek and river margins. DISPERSAL

ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults havebeen found in April – November. BEHAVIOR: Nocturnal, adults take cover during the day inburrows 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 1835LONG 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, andNovember – 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, UASMcollections

Omophron ovale G. Horn 1870OVAL ROUND SAND BEETLE

Omophron ovale G. Horn, 1870:75Omophron concinnum Casey, 1897:302Omophron gemma Casey, 1897:304Omophron 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,


ponds, and reservoirs. DISPERSAL ABILITIES: Macropterous, probably capable of flight. Strongburrower. SEASONAL OCCURRENCE: Adults have been found in April – October; copulatingpairs in July; teneral adults in July and October. BEHAVIOR: Nocturnal, adults take cover duringthe 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 1870ROBUST ROUND SAND BEETLE

Omophron robustus G. Horn, 1870:73Omophron brevipenne Casey, 1909:276Omophron 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, andbrooks. MICROHABITAT: On open wet, sandy, sometimes alkaline ground. DISPERSAL ABILITIES:Macropterous, probably capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults havebeen found in May – September; teneral adults in August. BEHAVIOR: Nocturnal, adults takecover 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 1897SOLID ROUND SAND BEETLE

Omophron solidus Casey, 1897:303Omophron lawrencei Hatch, 1953:69

Distribution: Native, New World. USA – CA, OR.Way of Life: MACROHABITAT: Lowlands to midlands, 119 – 1052 meters altitude, beaches ofwater bodies. MICROHABITAT: Moist sandy ground near water. DISPERSAL ABILITIES:Macropterous, probably capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adultshave been found in April – November. BEHAVIOR: Nocturnal, adults take cover during the dayin burrows dug in the ground. Adults are gregarious.References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections

Omophron sphaericum Chevrolat 1835ROUND 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.)


Omophron tessellatum Say 1823MOSAIC ROUND SAND BEETLE

Omophron tessellatus Say, 1823:152Omophron Lecontei Dejean, 1831:582Omophron tesselatus LeConte, 1848:447Omophron 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 oflakes, 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 thesunshine, usually adults take cover during the day in burrows dug in the ground (adult andlarva). Adults are gregarious and often attracted to lights at night.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS,NMNH collections

LORICERITAELoricerini LeConte 1861

SPRINGTAIL HUNTERS

Loricerini LeConte, 1861:6

Number of worldwide genera: One.Number of Western Hemisphere genera: One.Taxonomy: Stable. Adelphotaxon: Elaphrini Latreille 1802Geographic status: Holarctic, Neotropical.References: Ball & Erwin (1969), Erwin (2001)

Loricera Latreille 1802SPRINGTAIL HUNTERS

(Plate 12)

Loricera Latreille, 1802:88Lorocera 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).


Habitat: According to Lindroth (1961), adults of this genus are very hygrophilous and veryactive in the sunshine, running and flying. Adults are predaceous on springtails (Collembola), asare the larvae that possess adhesive mouthparts. Adults use their stiff long setae on the basalsegments of their antennae as an enclosing trap to capture springtails. Larvae use the apicalpart of their galea as a sticky trap to capture springtails. Adults hibernate.References: Ball & Erwin (1969), Erwin (2001), Lindroth (1961)

Loricera aptena Ball & Erwin 1969WINGLESS 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 capableof 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 183310-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 – 585meters altitude. MICROHABITAT: Adults of this species occur near fresh water (e.g. borders ofponds, lakes, marshes, slow rivers and brooks, and seepages), but they are not as hygrophilousas L. pilicornis adults; they also are often found among grass and moss in the open some distancefrom 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 alsorun 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 1851PITTED 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


are not as hygrophilous as L. pilicornis adults; they are often under stones, boards, and fallentrees on clay-mixed sandy ground up slope from the water’s edge; clay ground at edges ofvernal 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, butalso 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 1853BERING 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 partiallyshaded rich organic ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight.SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: See descriptionunder genus, above.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS,NMNH collections

Loricera pilicornis pilicornis (Fabricius) 1775HAIRY-HORNED SPRINGTAIL-HUNTER

Carabus pilicornis Fabricius, 1775:243Carabus seticornis O.F. Müller, 1776:79Loricera vinearum Geoffroy, 1785:43Loricera aenea Latreille, 1806:224Loricera semipunctata Eschscholtz, 1833:25Loricera alpina Heer, 1838:44Loricera rufilabris Motschulsky, 1845:340Loricera californica LeConte, 1863:3Loricera neoscotica LeConte, 1863:3Loricera uteana Casey, 1920:147Loricera sierrae Van Dyke, 1925:113Loricera apennina Binaghi, 1942:135Loricera 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 organicdebris, covered with sparse vegetation and shaded by shrubs and tall plants at the borders ofeutrophic marshes, ponds, lakes, pools (often polluted by refuse), slow rivers, brooks, and


irrigation ditches; fens, swamps, Carex marshes, low forests, flood-plain forests, mud flats, wetmeadows and pastures, cultivated fields (e.g., barley, dry bean). Also, found in apple orchards inBC 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 inlake-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 adultsin June – September. BEHAVIOR: Diurnal and nocturnal, active in the sunshine or adults takecover 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 andswamps, 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 standingtree stumps, and in the ground (as deep as 20 cm). Adults are predaceous on springtails andinsect 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 1863ROUND-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: MACROHABITAT: Midlands to mountains, 930 – 2928 meters altitude.MICROHABITAT: Adults are found on higher, dry ground in cloud, oak, oak-pine forests withArbutus, 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; macropterousindividuals found only in large populations. SEASONAL OCCURRENCE: Adults have been foundin January, March, and May – December. Larvae found in August. BEHAVIOR: See descriptionunder genus, above. Adults take cover in the day under fallen trees and in the leaf litter. Larvaeand pupae also found under large embedded stones.References: Ball & Erwin (1969), Erwin (2001), Data from CAS, NMNH, UASM collections

CARABITAECarabini 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 1834Geographic status: All Realms.References: Erwin (2001), Lorenz (2005)


Carabina Latreille 1802SEARCHERS

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 globalsynthesis. Adelphotaxon: Ceroglossina Vacher de Lapouge 1927Geographic status: Holarctic, Neotropical, Neaustral, Oriental, Australasian, AfrotropicalReferences: Erwin (2001), Lorenz (2005)Notes on the Calasoma-group: Gidaspow (1959) summarizes the feeding habits of theCalasoma-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. Theyusually 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 ofconsiderable size. The larvae eat the liquid and fatty parts of the caterpillars and often leave themeal before it is finished, thus destroying more caterpillars than they actually need. The larvaefavor the pupae of butterflies, except those that are hidden in cocoons. However, there arerecords that cocoons of Malacosoma americana and Malacosoma disstria have been destroyed. Usuallya larva makes a hole on the side of the pupa, between the segments, enlarges it, and then pushesthe front part of its body inside. It leaves the pupa when there is virtually nothing left but ashell. While eating, the larva is well hidden beneath the old pupa cases. The entrance hole madeby 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 thatamong the pupae killed by the larvae of Calosoma sycophanta 75 per cent were females. One larvaof Calosoma as large as sycophanta or scrutator is able to kill about 50 caterpillars or 15 pupaebetween the time of hatching and pupation. The adults of Calosoma seize the caterpillars in themiddle 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 softabdomen, such as some species of the family Chrysomelidae, it is attacked from below. Incaptivity, the beetles can also eat beef meat, but after a week or so, they refuse to feed, andcaterpillars must be given to them. Apparently, the species of Calosoma are not affected by thediseases from which some caterpillars often suffer. Experiments showed that Calosoma sycophantawas not affected by a disease of the gypsy moth known as “wilt.” Cannibalism among thelarvae is not uncommon and, to a certain extent, reduces the number of larvae even when foodis abundant. The best known species of the genus, Calosoma sycophanta, is, as is said above, moreor less adapted to the life of the gypsy moth and is very active when the caterpillars of thegypsy moth are present. Most of the larvae hatch during the pupal stage of the moth. In spiteof the fact that sycophanta attacks other caterpillars and pupae, it is never numerous in theregions where the gypsy moth is absent. Both larvae and adults are very good climbers; theymass on trunks of the trees under branches where the pupae of the gypsy moth are usuallyhidden and devour them on the spot. Occasionally they attack the imago also, mostly a femalelaying eggs. Calosoma frigidum lives chiefly on Heterocampa guttivitta and other Lepidoptera thatpupate on the ground. The larvae of frigidum are not good climbers and feed on pupae that areusually hidden in cells under leaf mold. Calosoma calidum is also more terrestrial in habits; the


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 alsoadapted 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 thatthey are not such good climbers as is Calosoma sycophanta. Calosoma externum is known to feed onlocusts and army worms.”

Callisthenes Fischer von Waldheim 1820BEAUTIFUL BLACK SEARCHERS

(Plate 13)

Callisthenes Fischer von Waldheim, 1820: pl. 7Chrysostigma Kirby, 1837:19 [valid subgenus for native Nearctic and Neotropical species]Tapinosthenes Kolbe, 1895:56Microcallisthenes Apfelbeck, 1918:161Callistenia Vacher de Lapouge, 1929:2 [valid subgenus for native Nearctic species]Lyperostenia Vacher de Lapouge, 1929:3See Lorenz (2005) for additional Old World subgenera.

Number of worldwide species: 53.Number of Western Hemisphere species: 25.Taxonomy: Stable. Adelphotaxon: Calosoma Weber 1801Type species: Callisthenes panderi Fischer von Waldheim, 1820Distribution: 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 arefully winged and fly, the latter also are climbers. The life history of many species in NorthAmerica is known (Burgess & Collins, 1917). Adults and larvae are ravenous carnivores,specializing upon caterpillars and pupae of Lepidoptera, never, as far as is known, monophagouslyrestricted to a single host.References: Erwin (2001), Gidaspow (1959), Lindroth (1961)

Callisthenes (Chrysostigma) affinis (Chaudoir) 1843RELATED BEAUTIFUL BLACK SEARCHER

Calosoma affine Chaudoir, 1843:746Calosoma triste LeConte, 1845:201Calosoma 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 sandyground with sparse vegetation. DISPERSAL ABILITIES: Macropterous, probably capable of flight.Moderate runners, and climbers on plants. SEASONAL OCCURRENCE: Adults have been found inApril – October. Females with eggs in July – August. BEHAVIOR: Adults are gregarious (sometimes


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) 1891LARGE 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: Likelynear wet or damp areas with shrubby vegetation or small trees in the Chihuahuan Desert andpossibly at the margins of cultivated lands. DISPERSAL ABILITIES: Macropterous, probably capableof 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) 1775FIERY HUNTER

Carabus calidus Fabricius, 1775:237Calosoma calida expansa Casey, 1897:344Calosoma calida laticollis Casey, 1897:344Calosoma calida stellata Casey, 1897:344Calosoma comes Casey, 1920:156Calosoma concreta Casey, 1920:157Chrysostigma 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, WYWay 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 ofopen 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 latespring 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, butwill 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, andthey will also eat tent caterpillars and army worms (Burgess and Collins, 1917). They can becollected by tying burlap sacks around the lower trunks of trees, a device that congregates theirprey; they will also come to sugar used for “sugaring” for moths. They are both nocturnal, andaccording to Larochelle (1975), diurnal, as well. Only one specimen was collected on PlummersIsland in Maryland, in 1913 “at light.” There have been no recent records. Hibernation takes placein the soil up to 6 cm deep. A long and detailed report on their bionomics is given by Burgess andCollins (1917) and is briefly summarized in Larochelle & Larivière (2003).


References: Erwin (2001), Burgess and Collins (1917), Larochelle (1975), Larochelle & Larivière(2003), Lindroth (1961), Data from NMNH collection

Callisthenes (Chrysostigma) cancellatus (Eschscholtz) 1833LATTICED BEAUTIFUL BLACK SEARCHER

Calosoma cancellatum Eschscholtz, 1833:23Calosoma aenescens LeConte, 1854:16Calosoma esuriens Casey, 1913:64Calosoma transversa Casey, 1913:65Calosoma sagax Casey, 1920:158Calosoma rectilatera Casey, 1920:158Calosoma 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 tulemarshes in CA. DISPERSAL ABILITIES: Adults are submacropterous, probably incapable of flightand are moderate runners and occasional climbers (on shrubs). SEASONAL OCCURRENCE: Adultsare active in March – September; females with eggs in May – June. Larvae are present in thefauna in summer and thus over winter. BEHAVIOR: Adults are diurnal; active in the sunshine onthe 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 feedupon larvae of the moth Peridroma saucia (Hübner).References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNHcollection

Callisthenes (Callistenia) dawsoni Dajoz 1997DAWSON’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 openground which is covered almost everywhere with calcium salts and with sparse vegetationcover dominated by Graminea and some Artemisia shrubs. DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave been found in August. BEHAVIOR: Adults are predaceous on caterpllars.References: Dajoz (1997), Erwin (2001)

Callisthenes (Callistenia) dietzi (Schaeffer) 1904DIETZ’S BEAUTIFUL BLACK SEARCHER

Calosoma Dietzii Schaeffer, 1904:197Callisthenes gravidulus Casey, 1913:68

Distribution: Native, New World. USA – CA.


Way of Life: MACROHABITAT: Midlands to mountains, 800 – 2700 meters altitude.MICROHABITAT: Open dry grasslands and mountain meadows. DISPERSAL 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) 1857DIFFERENT BEAUTIFUL BLACK SEARCHER

Calosoma discors LeConte, 1857:31Callisthenes 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 grasslandsand 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 NMNHcollection

Callisthenes (Callistenia) lariversi (Van Dyke) 1943LARIVERS’ 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 vagilitylimited 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) 1870BROAD BEAUTIFUL BLACK SEARCHER

Calosoma latipenne G. Horn, 1870:70Calosoma arcuata Casey, 1897:343Callisthenes tularensis Casey, 1913:68Callisthenes optimus Casey, 1913:69

Distribution: Native, New World. USA – CA, NV.Way of Life: MACROHABITAT: Lowlands to midlands, 113 – 1300 meters altitude, in MojaveDesert vegetation. MICROHABITAT: Sandy soils with desert scrub in lowlands; sandy soils androcky 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


Callisthenes (Chrysostigma) lepidus (LeConte) 1845NEAT 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: Sub-brachypterous, 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 NMNHcollection

Callisthenes (Callistenia) luxatus (Say) 1823DISLOCATED BEAUTIFUL BLACK SEARCHER

Calosoma luxata Say, 1823:149Carabus Zimmermanni LeConte, 1848:345Calosoma striatulum LeConte, 1859:4Calosoma pimelioides Walker, 1866:312Callisthenes luxatus opacus Géhin, 1885:70Callisthenes exaratus Casey, 1913:72Callisthenes tegulatus Casey, 1913:72Callisthenes tegulatus viator Casey, 1913:72Callisthenes pustulosus Casey, 1913:73Callisthenes diffractus Casey, 1913:75Callisthenes reflexus Casey, 1920:164Callisthenes utensis Casey, 1920:165Callisthenes semotus Casey, 1920:166Callisthenes debilis Casey, 1920:167Callisthenes parowanus Casey, 1920:167Calosoma luxatum zimmermanni striatus Breuning 1928:86Calosoma 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 withindry forests (e.g., Pinus). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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), Datafrom CAS, NMNH collections


Callisthenes (Callistenia) moniliatus LeConte 1852COLLARED BEAUTIFUL BLACK SEARCHER

Callisthenes moniliatus LeConte, 1852:200Calosoma laqueatum LeConte, 1860:318Carabus bicolor Walker, 1866:313Callisthenes concinnus Casey, 1913:66Carabus 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 openground 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 NMNHcollection

Callisthenes (Callistenia) monticola (Casey) 1897MOUNTAIN BEAUTIFUL BLACK SEARCHER

Calosoma monticola Casey, 1897:342Callisthenes 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: Opendry ground in sagebrush areas (Artemisia sp.). DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary – 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) 1885MORRISON’S BEAUTIFUL BLACK SEARCHER

Calosoma Morrisonii G. Horn, 1885:128Calosoma 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: Opendry 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


Callisthenes (Chrysostigma) obsoletus (Say) 1823OLD BEAUTIFUL BLACK SEARCHER

Calosoma obsoleta Say, 1823:149Calosoma indistinctum LeConte, 1845:208Calosoma 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, WYWay 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 favoredby 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 dungand dead prostrate weeds; sometimes active by the hundreds on cloudy days. Adults foundoverwintering 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 willeat 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 1959OREGON 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 1959PLACER 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


Callisthenes (Callistenia) schaefferi (Breuning) 1928SCHAEFFER’S BEAUTIFUL BLACK SEARCHER

Calosoma irregulare Schaeffer, 1915:235Calosoma discors schaefferi Breuning, 1928:79Calosoma 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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April– July. BEHAVIOR: Unknown.References: Erwin (2001), Data from CAS, NMNH collectionsNote: Gidaspow (1959) states that she did not see Hatch’s type, nor could she make much ofthe description. I have compared the Holotype of C. striatus (now in NMNH) with Scheaffer’stype of C. irregulare (also in NMNH) and they represent the same species.

Callisthenes (Chrysostigma) semilaevis (LeConte) 1852SEMI-SMOOTH BEAUTIFUL BLACK SEARCHER

Calosoma semilaeve LeConte, 1852:199Calosoma semilaeve davidsoni Casey, 1914:33Calosoma 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 pea-vines. Markedly favored by human activities. DISPERSAL ABILITIES: Macropterous, probably capableof flight. Adults and larvae are occasional climbers. SEASONAL OCCURRENCE: Adults have beenfound 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 HyEdwards), tenebrionids, elaterids, and carabids. Adults are most active in April – May.References: Erwin (2001), Burgess & Collins (1917), Larochelle & Larivière (2003), Data fromNMNH collection

Callisthenes (Chrysostigma) simplex (LeConte) 1878SIMPLE 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 collectionNotes: 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.


Callisthenes (Callistenia) subaeneus (Chaudoir) 1869COPPERY 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) 1915ROUGH BEAUTIFUL BLACK SEARCHER

Calosoma subasperatum Schaeffer, 1915:235Callisthenes 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) 1852LUKEWARM BEAUTIFUL BLACK SEARCHER

Calosoma tepidum LeConte, 1852:199Calosoma irregulare Walker, 1866:312Calosoma tepida caelator Casey, 1913:61Calosoma tepida indigens Casey, 1913:61Calosoma pellax Casey, 1920:160Calosoma semicuprea Casey, 1920:161Calosoma 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 lowvegetation in cultivated fields (e.g, pea), rangelands, and along roadsides. DISPERSAL

ABILITIES: Submacropterous, probably incapable of flight. SEASONAL OCCURRENCE: Adultshave been found in May – August; teneral adults in May. BEHAVIOR: Adult are predaceouson caterpillars and grasshoppers.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNHcollection


Callisthenes (Callistenia) wilkesii LeConte 1852WILKES’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 meadowssometimes surrounded by woodlands. MICROHABITAT: Adults are found on dry ground in theopen with low vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagilitylimited 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 1880BEAUTIFUL THICKISH SEARCHERS

(Plate 15)

Calopachys Haury, 1880:164Eutelodontus 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, 1880Distribution: Neotropical (4 species). Although far enough south to be Neotropical, all of thespecies 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) 1846HURTFUL BEAUTIFUL THICKISH SEARCHER

Calosoma blaptoides Putzeys, 1846:400Eutelodontus aztecus Géhin, 1881:cxxxiiEutelodontus 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-pineforests and meadows. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave been found in May – August. BEHAVIOR: Nocturnal, adults take cover under stones and inleaf litter in the day.References: Erwin (2001), Gidaspow (1959), Data from CAS, UASM collections


Calopachys bohnei (Korell & Frisch) 2004BOHNE’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: Underfallen branches and fallen trees on rocky soils of volcanic origin beneath fir-oak forest (Abiesguatemalensis and Quercus magnolifolia Née, Q. castanea A. DC.). DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave been found in July. BEHAVIOR: Unknown.References: Korell & Frisch (2004), Data from NMNH collectionNotes: This and the following species seem to be altitudinally separated, however both speciesare represented (NMNH) in a series collected in July by H.H. Smith at “Omiltemi, 8000 ft.”

Calopachys omiltemius (Bates) 1891OMILTEMI 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 montanerain and cloud forests, and subtropical mixed oak-pine forests. MICROHABITAT: On rocky soilsof volcanic origin beneath forests of Quercus magnolifolia Née, Q. castanea A. DC., with someindividual 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) 1880GREENISH 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 largestones in oak forest. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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 collectionNotes: Only two specimens are known of this species.


Calosoma Weber 1801CATERPILLER HUNTERS

(Plates 14 and 16)

Calosoma Weber, 1801:20Callitropa 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, 1866:307Calamata Motschulsky, 1866:307Camedula Motschulsky, 1866:304Blaptosoma 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, 1927:146Paratropa Vacher de Lapouge, 1929:3Acampalita Vacher de Lapouge, 1929:9Catastriga Vacher de Lapouge, 1929:9Caludema Jeannel, 1940:68

Number of worldwide species: 84.Number of Western Hemisphere species: 55.Taxonomy: Stable. Adelphotaxon: Callisthenes Fischer von Waldheim 1820Type species: Carabus sycophanta Linné, 1758Distribution: Holaractic, Neotropical, Neaustral, Oriental, Australasian, Afrotropical, Native,New World (54 species), Nearctic (13 species), Introduced to the Nearctic from Europe (onespecies), 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 isknown (Burgess & Collins, 1917). Adults and larvae are ravenous carnivores, specializing uponcatepillars and pupae of Lepidoptera, never, as far as is known, monophagously restricted to asingle 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 attractedto lights. Adults of some species, however, have reduced wings and do not fly; however, theseare 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 1869SHORT 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 coveredplains, in cultivated land (e.g. cotton fields), and in sandy areas between the coast and oak-covered foothills in Colombia. MICROHABITAT: On partly shaded sandy ground. DISPERSAL

ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have beenfound in January – April, and November – December. BEHAVIOR: Unknown.References: Erwin (1991, 2001), Gidaspow (1963), Data from CAS, NMNH collections


Calosoma (Castrida) alternans (Fabricius) 1792RECIPROCAL CATERPILLER HUNTER

Carabus alternans Fabricius, 1792:146Calosoma Laterale Dejean, 1826:199Calosoma granulatum Perty 1830:9Calosoma imbricatum Brullé, 1837:42Calosoma Brulléi Blanchard, 1843:42Calosoma armatum Reiche, 1842:377Callistriga coxale Motschulsky, 1865:307Calamata rugata Motschulsky, 1865:308Calosoma 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) andSpodoptera 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 collectionNotes: In NMNH, there is a male specimen labeled Mali, W. Africa, CICP Project, August,1987, “Nara, Acacia.”

Calosoma (Carabomimus) altipeta (Jeannel) 1940VOLCANO 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 flightlessthus 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 1834ANGULATE CATERPILLER HUNTER

Calosoma angulatum Chevrolat, 1834: no. 44Calosoma angulicolle Chaudoir, 1869:377Calosoma uniforme Géhin, 1885:63Calosoma Forreri Géhin, 1885:64


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, premontanemoist forest, thorn scrub, oak-savannah, possibly in cultivated land. MICROHABITAT: Leaf litterin 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 1831BLACK CATERPILLER HUNTER

Calosoma Anthracinum Dejean, 1831:569Calosoma rufinum Géhin, 1885:66

Distribution: Native, New World. MÉXICO – GO.Way of Life: MACROHABITAT: Uplands, 1828 meters altitude, in montane forests of diverseand complex subtropical mixed oak-pine forests. MICROHABITAT: On the ground. DISPERSAL


OCCURRENCE: Unknown. BEHAVIOR: Unknown.References: Erwin (2001), Gidaspow (1959), Data from NMNH collection

Calosoma (Blaptosoma) anthracinum microgonum Bates 1891SMALL-ANGLED CATERPILLER HUNTER

Calosoma laeve microgonum Bates, 1891:225

Distribution: Native, New World. MÉXICO – DF, JA, PU, VC.Way of Life: MACROHABITAT: Uplands to mountains, 1500 – 2520 meters altitude.MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagilitylimited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown.References: Erwin (2001), Gidaspow (1959), Data from NMNH collection

Calosoma (Castrida) argentinensis Csiki 1927ARGENTINE CATERPILLER HUNTER

Calosoma argentinensis Csiki, 1927:11Calosoma Antiquum Dejean, 1831:561Calosoma belsaki Dallas, 1939:782Calosoma cuprifulgens Dallas, 1939:784Calosoma miniaturum Dallas, 1939:784Calosoma ballasi Dallas, 1939:788Calosoma pampeanum Belsak, 1939:788Calosoma dallasi Dallas, 1939:789


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 thedrier regions of its range. MICROHABITAT: On the ground in xeophytic thickets and Chacoforests, dry valleys of the pampas, and in desert scrub and grassy areas on barren soil; also inpastures and cultivated land (mostly alfalfa fields). Found in great numbers on the open saltflats 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 predaceouson insect larvae, principally caterpillars of Colias lesbia (F.). Adults appear newly emerged in lateOctober, 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) 1940ROUGH 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: Adultshave been found in May – August, and November. BEHAVIOR: Adults take cover under barkand fallen trees of conifers during the day.References: Erwin (2001), Gidaspow (1959), Data from NMNH collection

Calosoma (Blaptosoma) atrovirens atrovirens Chaudoir 1869BLACKISH-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: Oakforests. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. 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 1891BROAD-NECKED CATERPILLER HUNTER

Calosoma explanaticolle Bates, 1891:225Calosoma obscurum Géhin, 1885:65

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


Way of Life: MACROHABITAT: Midlands to mountains, 1040 – 2408 meters altitude, in tropicaldeciduous 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 1850GOLD-RINGED CATERPILLER HUNTER

Calosoma aurocinctum Chaudoir, 1850:420Calosoma 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 collectionsNotes: In NMNH, there are two specimens that may be mislabeled, one labeled Chile and theother “Tucuman” Argentina.

Calosoma (Carabomimus) bolivianum Géhin 1885BOLIVIAN 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 1869BRIDGES’ 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 vagilitylimited to walking or running. SEASONAL OCCURRENCE: An adult was found in October.BEHAVIOR: Unknown.References: Erwin (2001), Gidaspow (1963), Data from NMNH collection


Calosoma (Carabomimus) bulleri Breuning 1943BULLER’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 1959CHIHUAHUA 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-oakwoodlands, MICROHABITAT: In meadows and on lake margins. DISPERSAL 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 1869SCARRED CATERPILLER HUNTER

Calosoma cicatricosum Chaudoir, 1869:374Calosoma balli, Rotger, 1976:294

Distribution: Native, New World. MÉXICO – DF, MX.Way of Life: MACROHABITAT: Uplands to mountains, 2166 – 2468 meters altitude.MICROHABITAT: DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 1869RIBBED 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 walkingor running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown.References: Erwin (2001), Gidaspow (1959), Data from NMNH collections


Calosoma (Carabomimus) depressicolle Chaudoir 1869FLAT-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 leaflitter or among grasses. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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) 1924DIGUET’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/pineforest, scrub forest. MICROHABITAT: Found in clearings. DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave 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 1891SMALL CATERPILLER HUNTER

Calosoma diminutum Bates, 1891:227Callisthenes laevissimum Casey, 1920:170

Distribution: Native, New World. MÉXICO – MX, VC.Way of Life: MACROHABITAT: Uplands to mountains, 1733 – 2750 meters altitude.MICROHABITAT: Pastures on slopes. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility 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 1910SOLITARY CATERPILLER HUNTER

Calosoma eremicola Fall, 1910:91Calosoma rugosipennis Schaeffer, 1911:113Calosoma hospes Casey, 1913:63Acamegonia peregrinatrix incerta Vacher de Lapouge, 1924:38

Distribution: Native, New World. MÉXICO – BJ; USA – CA (+ SAN CLEMENTE AND SANTA

CATALINA ISLANDS), NM, NV.


Way of Life: MACROHABITAT: Lowlands to uplands, 100 – 1600 meters altitude, in EasternMojave Desert vegetation zone. MICROHABITAT: Found in high desert, as well as in fields andalong 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 NMNHcollection

Calosoma (Callitropa) externum (Say) 1823CHURCHYARD CATERPILLER HUNTER

Carabus externum Say, 1823:150Calosoma 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: Adultshave been found in February, April – October, and December. BEHAVIOR: Adults are oftenfound at lights at night and occasionally in drift material at the margin of water bodies; they arepredaceous 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 1884FLOHR’S CATERPILLER HUNTER

Carabomimus Flohri Bates, 1884:262Calosoma 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 theground among grasses and in leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inMay 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 1837COLD-COUNTRY CATERPILLER HUNTER

Calosoma frigidum Kirby, 1837:19Calosoma 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.


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, orconiferous forests, tree plantations (e.g., Pinus), and orchards. DISPERSAL ABILITIES: Macropterous,capable of flight. Frequent climber on trees and shrubs. SEASONAL OCCURRENCE: Adults havebeen found in March, and May – November; copulating pairs in May – June; gravid females inMay – June; tenerals in May – September. BEHAVIOR: Diurnal and nocturnal, adults are activeon trees (e.g., Abies, Picea, Populus, Quercus) and shrubs, or adults take cover under the loose barkof dead trees, in leaf litter, and under fallen trees. Adults are gregarious. Eggs are laid singly inthe 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 cmdeep on average) and in rotten tree-stumps. Life span can be more than 2 years. Adult arepredaceous on lepidopterous caterpillars, cicadas, and locusts. Adults and perhaps larvae mayplay 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 presenceof large numbers of forest defoliators. Adults occasionally found at lights at night. Frequentflier, often swarming during the day. Frequent in drift material.References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNHcollection

Calosoma (Castrida) fulgens Chaudoir 1869SHINING 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: Adultshave 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 1838GALÁPAGOS 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January and December. BEHAVIOR: Unknown.References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963),Data from NMNH collection


Calosoma (Carabomimus) gebieni (Breuning) 1928GEBIEN’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 1831SMOOTH CATERPILLER HUNTER

Calosoma Glabratum Dejean, 1831:565Carabosoma bolivianum Géhin, 1885:65

Distribution: Native, New World. BOLIVIA; COLOMBIA; PANAMÁ; PERÚ.Way of Life: MACROHABITAT: Midlands to mountains, 730 – 2650 meters altitude.MICROHABITAT: In grasslands and cultivated fields. DISPERSAL ABILITIES: Macropterous, probablycapable 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 1953:10DARWIN’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 onvolcanic soils. DISPERSAL ABILITIES: Brachypterous, wings slightly reduced, hence flightless thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inAugust – September. BEHAVIOR: Unknown.References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963),Data from NMNH collection

Calosoma (Castrida) granatense floreanum (Basilewsky) 1968FLOREANA 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 speciesis 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 beenfound 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


Calosoma (Castrida) granatense granatense Géhin 1885WIDE-LINES CATERPILLER HUNTER

Calosoma granatense Géhin, 1885:59Calosoma galapagoum Howard, 1889:191Calosoma Howardi Linell, 1898:251Calosoma 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 aridand lower transition vegetation belt, and lower moist forest zone. MICROHABITAT: Desender etal. (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 aridand lower transition vegetation belts at the lower elevations of the volcanoes. On Fernandinaand on all separate volcanoes of Isabela the species also occurs at higher elevations, mostly inso-called “pampa” or meadow vegetation. Our morphometric data show that in this species thehindwings vary in size and shape (cfr. Fig. 2). As far as we know this is the first case of wingpolymorphism reported for a beetle species in the Galapagos. Full-winged specimens, whichare more numerous in lowland populations, mostly have functional in direct flight muscles andindeed can exhibit considerable flight activity during nights after heavy rainfall. In lowland dryarid and transition scrub vegetation, this species most probably feeds especially on lepidopteralarvae (as deduced from our few field observations of feeding beetles) and we hypothesize thatthe adaptive significance of light activity is mainly related to the search for new sites or treeswith abundant food. An interesting case of variation in dispersal power between populationsof Calosoma granatense is observed on Volcan Cerro Azul (cfr. Fig. 3), one of the highest volcanoesin the entire archipelago, reaching 1700m altitude and moreover showing an inversion in thepresence of different vegetation types along its slopes (cfr. HAMANN, 1981). Because of itsheight, 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 arelatively luxuriant vegetation on nearly its complete southwestern slopes, at least during theannual rainy season. Near the coast there is a dry deciduous steppe forest, which from about250m 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 isindicated by a somewhat dryer and more open grassland vegetation with a small shrub speciesbecoming more common. This zone indicates the mean upper level of clouds and is succeededby an increasingly xerophytic and shrubby vegetation higher up the volcano. From the inversionzone onwards to the top there is thus again a more patchy and shrubby vegetation, characterizedas a steppe scrub. The inversion in vegetation is also indicated by several plant species, such asOpuntia 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. Calosomagranatense beetles were collected by us during 1986 along almost the entire slope of this volcanoup 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 mediumaltitude and higher again in the steppe scrub athigher altitute. The results of an Analysys of


Variance (F = 15.75,p < 0.001, n=124) between the wing development in these differentpopulations are highly significant,whereas a posteriori multiple range tests indicate the occurrenceof three main groups of populations coinciding with the three major types of vegetation. Wehypothesizethat wing development has been reduced at medium altitude following pastemigration events by full winged ancestral individuals, because there is probably relaxed or noselection pressure acting in this open habitat to retain a high dispersal power: feeding only cantake place at the soil surface level in the continuous and low vegetation (in space as well as intime); at lower altitudes as well as above the inversion zone the vegetation structure is threedimensional and complex, composed of trees and scrubs which are moreover patchily distributed;in these circumstances there is probably a continuous selection pressure promoting the retentionof a higher dispersal ability. The somewhat lessened wing development of beetles from the topzone could then be the inevitable result of the higher possibility for unidirectional gene flow tooccur 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 anadaptive significance counteracting the effects of losses due to gene flow from an area ofrelaxed selection.” DISPERSAL ABILITIES: Wing polymorphic, SEASONAL OCCURRENCE: Adultshave been found in January – May, and September – November. BEHAVIOR: Fully-wingedadults occasionally found at lights at night.References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963),Data from NMNH collectionNotes: There are old yet doubtful records of this species occurring in Colombia and Perú.

Calosoma (Blaptosoma) haydeni haydeni G. Horn 1870HAYDEN’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; sometimesin 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 midAugust; teneral adults in July. BEHAVIOR: Predatory on caterpillars and pupae at night; adultstake 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 1891HOLEY-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-oakwoodlands, grasslands. 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, and September – November. BEHAVIOR: Nocturnal, adultstake cover in the day under stones.References: Erwin (2001), Gidaspow (1959), Data from CAS, NMNH, UASM collections

Calosoma (Blaptosoma) laeve Dejean 1826DEJEAN’S CATERPILLER HUNTER

Calosoma Laeve Dejean, 1826:210Calosoma 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 groundnear water. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. 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 1928HÖGE’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 nearwater. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. 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 1869HÖGE’S SMOOTH CATERPILLER HUNTER

Calosoma laevigatum Chaudoir, 1869:374Calosoma 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 orrunning. 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) 1968LELEUPS’ CATERPILLER HUNTER

Castrida leleuporum Basilewsky, 1968:200

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


Way of Life: MACROHABITAT: Midlands, 500 – 750 meters altitude, island humid fern-sedgeand Sphagnum vegetation. MICROHABITAT: On moist soil beneath Miconia shrubs in humidgrasslands and in Sphagnum bogs. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary, April, and December. BEHAVIOR: Adults hibernate deep in the soil on damp steepslopes, 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 1925LINELL’S CATERPILLER HUNTER

Calosoma Linelli Mutchler, 1925:221Calosoma 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 openpampa habitats with some shrubs. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary. BEHAVIOR: Unknown.References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963),Data from NMNH collection

Calosoma (Callitropa) macra (LeConte) 1853LONG CATERPILLER HUNTER

Callitropa macra LeConte, 1853:400

Distribution: Native, New World. MÉXICO – CL, TM; USA – AR, LA, NM, OK, TX. Wayof Life: MACROHABITAT: Midlands, 1000 – 1600 meters altitude, in open country. MICROHABI-TAT: Unknown. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE:Adults have been found in February – November. BEHAVIOR: Adults are occasionally foundat lights at night.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

Calosoma (Carabosoma) marginale Casey 1897RIMMED CATERPILLER HUNTER

Calosoma marginalis Casey, 1897:340Calosoma lugubre LeConte, 1853:400Calosoma 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:


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

Calosoma (Carabomimus) morelianum Bates 1891MORELOS 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) 1891ORIZABA 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 1876PALMER’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, ondesert island(s) off the coast of México. MICROHABITAT: Rocky pine-oak savannah (Pinus radiateD. 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 becauseof over-grazing by introduced and now feral goats. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January, and March – April. BEHAVIOR: One specimen was found at the very summitof the Island.References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collectionsNote: A single specimen in CAS is labeled Isla San Benito, West Island.


Calosoma (Carabosoma) parvicolle Fall 1910SMALL-NECKED CATERPILLER HUNTER

Calosoma parvicollis Fall, 1910:90Calosoma prominens Bates, 1881:21Calosoma clemens Casey, 1914:32Calosoma 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 metersaltitude, on deserts and sand dunes, MICROHABITAT: Open dry ground with sparse vegetation.DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults havebeen found in March – July, and September. BEHAVIOR: Adults are nocturnal and diurnal,gregarious, and predaceous on caterpillars, orthopterans, and spiders. Occasionally adults foundflying to lights at night.References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNHcollection

Calosoma (Carabosoma) peregrinator Guérin-Méneville 1844TRAVELING CATERPILLER HUNTER

Calosoma peregrinator Guérin-Méneville, 1844:255Calosoma carbonatum LeConte, 1862:53Calosoma affine Bates, 1881:21Calosoma peregrinator ingens Casey, 1913:62Calosoma peregrinator amplipennis Casey, 1913:62Calosoma subgracilis Casey, 1913:63Calosoma 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 thornscrub with yucca at riparian habitats. MICROHABITAT: On open ground covered with somevegetation 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 incavities in the ground. Adult are predaceous on caterpillars. Occasional climber on trees, adultstake cover under stones. Adults occasionally found at lights at night.References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH,UASM collectionsNotes: One female specimen in NMNH is labeled “Bogotá, Oct. 11, 1965.”


Calosoma (Castrida) peruviense Mandl 1971PERÚ 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 1869FURBISHED 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: Openground, among grasses and in leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inApril – May, July – August, and October – November. BEHAVIOR: Nocturnal, adults take coverin the day under stones.References: Erwin (2001), Gidaspow (1959), Jeannel (1940), Data from NMNH, UASMcollections

Calosoma (Blaptosoma) porosifrons Bates 1891POROUS-FRONTED CATERPILLER HUNTER

Calosoma porosifrons Bates, 1891:229Calosoma laevifrons Breuning, 1931:621

Distribution: Native, New World. MÉXICO – DU.Way of Life: MACROHABITAT: Uplands to mountains, 2150 – 2800 meters altitude, in oak/pineforest. MICROHABITAT: Riparian, this species occurs at El Salto, Durango. El Salto is an old, butstill 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 burnedmany times, but still supports a good growth of moderate sized Yellow pines (Pinus ponderosaDougl. 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 coveredwith 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


Calosoma (Carabosoma) prominens LeConte 1853PROJECTING CATERPILLER HUNTER

Calosoma angulatum LeConte, 1852:199Calosoma 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 1862ATTRACTIVE CATERPILLER HUNTER

Calosoma protractum LeConte, 1862:52Callitropa dolens Chaudoir, 1869:376Callitropa truncata Haury, 1885:64

Distribution: Native, New World. MÉXICO – DU, GJ, GO, JA, MH, ML, NA, PU, OA, SO;USA – AZWay of Life: MACROHABITAT: Midlands to uplands, 1150 – 1707 meters altitude, in acaciascrub. MICROHABITAT: On the ground in high desert where cactus meets oak and mesquitethickets. 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, UASMcollections

Calosoma (Castrida) retusum (Fabricius) 1801BLUNT-TOOTHED CATERPILLER HUNTER

Carabus retusum Fabricius, 1801:174Calosoma laterale Kirby, 1819:379Calosoma Bonariense Dejean, 1831:560Calosoma patagonense Hope, 1838:129Calosoma proseni Dallas, 1944:5Calosoma 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 drierregions 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.


References: Erwin (2001), Gidaspow (1963), Data from CAS, NMNH collections

Calosoma (Castrida) rufipenne Dejean 1831RED-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 thelomas, pampas, and in cultivated fields. MICROHABITAT: On the ground in grassy and desertscrub 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 1826SAY’S CATERPILLER HUNTER

Calosoma Sayi Dejean, 1826:198Calosoma armata Laporte de Castelnau, 1835:156Calosoma alternans Jacquelin du Val, 1857:19Calosoma sayi abdominale Géhin, 1885:58Calosoma sayi virginica Casey, 1897:344Calosoma 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, intropical dry and very dry forests, cactus/acacia desert, tropical moist forests, premontanewet forests, and wet forests. MICROHABITAT: Found on the ground in open areas in cultivatedlands (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 arenocturnal, taking cover in the day under wood. Larvae are mostly diurnal. Adults arepredaceous 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 andshrubs. 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


Calosoma (Calosoma) scrutator (Fabricius) 1775EXAMINOR’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 hardwoodforests and dry oak/pine forests. MICROHABITAT: On the ground or climbing plants. DISPERSAL

ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found inJanuary, and March – November. BEHAVIOR: Adults are predaceous on tent and noctuidcaterpillars. 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, UASMcollections

Calosoma (Calosoma) splendidum Dejean 1831SPLENDID 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: Inpalm forests in southern Texas. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL

OCCURRENCE: Adults have been found in April – July. BEHAVIOR: Adults occasionally found atlights at night.References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNHcollection

Calosoma (Carabosoma) sponsum Casey 1897BETROTHED CATERPILLER HUNTER

Calosoma sponsa Casey, 1897:340Calosoma 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 NMNHcollection


Calosoma (Carabosoma) striatipenne Chaudoir 1869FURROWED-WINGED CATERPILLER HUNTER

Calosoma striatipenne Chaudoir, 1869:373Calosoma 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 1835LINED CATERPILLER HUNTER

Calosoma striatulum Chevrolat, 1835: no. 165Calosoma 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, disturbedareas, and pastures. MICROHABITAT: On the ground along road edges, among grasses in fields,and in disturbed areas under debris. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in Mayand August. Behavior: Nocturnal, adults take cover under debris.References: Erwin (2001), Gidaspow (1959), Data from NMNH, UASM collections

Calosoma (Calosoma) sycophanta (Linné) 1758AGREEABLE CATERPILLER HUNTER

Carabus sycophanta Linné, 1758:414Calosoma azureum Letzner, 1850:95Calosoma cupreum Letzner, 1850:95Calosoma marginatum Letzner, 1850:95Calosoma nigrocyaneum Letzner, 1850:95Calosoma purpureoaureum Letzner, 1850:95Calosoma severum Chaudoir, 1850:422Calosoma rapax Motschulsky, 1866:309Calosoma smaragdynum G. Rossi, 1882:204Calosoma Habelmanni Schilsky, 1888:180Calosoma purpureipennis Reitter, 1891:257Calosoma corvinum Heller, 1897:312Calosoma prasinum Vacher de Lapouge, 1907:124Calosoma solinfectum Jännichen, 1914:298Callipara anthracinum Vacher de Lapouge, 1924:42Calosoma Lapougei Breuning, 1927:174Calosoma nigrosuturale Jacquet, 1930:76Calosoma nigroaeneum Polentz, 1937:7


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 shadedmoderately 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 (adultand 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 – 4years. Adult are predaceous on caterpillars and pupae (e.g., Lymantriidae, Thaumatopoeidae)and on coccinellids. Larvae are predaceous on caterpillars and pupae. Members of this speciesplay 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 byGypsy Moth outbreaks, human activities, and spring floods along large rivers. Adults frequentlyfound at lights at night particularly in the spring.References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNHcollection

Calosoma (Castrida) trapezipenne Chaudoir 1869DIM-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 westernplateau of Argentina, uplands of Mendoza and San Juan Provinces. MICROHABITAT: On theground in xerophyte forests of Córdoba and on the bare hills of Río Negro in dry brush anddesert 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 1831ROAMING 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 coldwinters. MICROHABITAT: On the ground in thickets and cultivated fields. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inJanuary – 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


Calosoma (Blaptosoma) viridisulcatum laetulum (Jeannel) 1940PLEASANT 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 metersaltitude. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJuly – September. BEHAVIOR: Unknown.References: Erwin (2001), Gidaspow (1959), Data from NMNH collection

Calosoma (Blaptosoma) viridisulcatum viridilucens Beheim & Breuning 1943BRIGHT 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 1863PLEASANT CATERPILLER HUNTER

Calosoma viridisulcatum Chaudoir, 1863:114Calosoma 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 1848WILCOX’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, adjacentgardens and cultivated fields (e.g., pea, cotton), orchards (e.g., plum, peach) and on trees incities. DISPERSAL ABILITIES: Macropterous, capable of flight. Frequent climber on trees andplants. SEASONAL OCCURRENCE: Adults have been found in January – December; copulating


pairs in June; gravid females in June. BEHAVIOR: Nocturnal and diurnal, active in the sunshineon trees, or adults take cover during the day under stones or downed trees. Adults foundoverwintering in cavities in the ground (2.5 – 15 cm deep). Adults are able to hibernate through2 winters and their life span is at least 2 – 3 years. Adult are predaceous on caterpillars andgrasshoppers. 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 1758WORM & SLUG HUNTERS

(Plate 17)

Carabus Linné, 1758:413Tachypus 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, 1887:7Tanaocarabus Reitter, 1896:135 [valid subgenus for native Nearctic species]Homoeocarabus Reitter, 1896:144 [valid subgenus for Holarctic species]Goniocarabus Reitter, 1896:150Tmesicarabus Reitter, 1896:159Aulonocarabus Reitter, 1896:192 [valid subgenus for Holarctic species]Diocarabus Reitter, 1896:185 [valid subgenus for native Nearctic species]Cancellocarabus Lutshnik, 1924:49Neocarabus Hatch, 1949:144See 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 globalsynthesis. Adelphotaxon: Ceroglossus Solier 1848Geographic status: Holarctic: Native, New World (11 species), Holarctic (one species),Introduced (six species)References: Erwin (2001), Lorenz (2005)

Carabus (Tachypus) auratus Linné 1761AURORA BOREALIS WORM & SLUG HUNTER

Carabus auratus Linné, 1761:219

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


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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April– July, and November. BEHAVIOR: Mostly diurnal, active in the sunshine. Adults foundoverwintering in the ground in Europe. Life span is several years, at least in captivity. Adult arepredaceous on earthworms, caterpillars, insects, molluscs, and they also feed on carrion andvegetal matter in Europe. They are spring breeders. Very effective as a colonist.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961, 1985), Data fromNMNH collectionNotes: This species is a late introduction from Europe and has become established in thecostal districts of New England and inland to Vermont.

Carabus (Tachypus) cancellatus Illiger 1798LATTICED 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 grasseyareas, as well as in forest clearings. DISPERSAL ABILITIES: North Carolina specimens in NMNHare brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL

OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Nocturnal predator, butalso diurnal during the reporductive period. Mating occurs mainly in May – June, teneral adultsemerge in July – August.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth, et al. (1985), Data fromNMNH collection

Carabus (Diocarabus) chaimssonis Fischer von Waldheim 1820FISCHER’S WORM & SLUG HUNTER

Carabus chaimssonis Fischer von Waldheim, 1820: pl. 7Carabus brachyderus Wiedemann, 1821:110Carabus Groenlandicus Dejean, 1831:554Carabus groenlandicus washingtoni Casey, 1920:155Carabus 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 withdiscontinuous vegetation (e.g., Empetrum), or on rich organic ground covered with leaf litter onmoraines 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


OCCURRENCE: Adults have been found in May – September; copulating pairs in July; teneraladults in August – September. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagilitylimited to walking or running. BEHAVIOR: Nocturnal, adults take cover during the day understones or among dead leaves. Adults found overwintering in their own pupal cavity. Coitus lastsat least 5 – 8 minutes.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

Carabus (Tanaocarabus) finitimus Haldeman 1852HALDEMAN’S WORM & SLUG HUNTER

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

Distribution: Native, New World. USA – KS, OK, TX.Way of Life: MACROHABITAT: Lowlands, 274 – 289 meters altitude, in bottomland forestsalong 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, adultstake 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 1882FORRER’S WORM & SLUG HUNTER

Carabus Forreri Bates, 1882:261Carabus 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-pineforests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJune – September, and November. BEHAVIOR: Predator. Nocturnal, adults take cover duringthe 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 2003WILL’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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July.


BEHAVIOR: Predator. Nocturnal, adults take cover during the day in under stones, fallen trees,or wood chips.References: Deuve (2003)

Carabus (Carabus) goryi Dejean 1831GORY’S WORM & SLUG HUNTER

Carabus limbatus Say, 1825:77Carabus Goryi Dejean, 1831:544Carabus 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 mositforests. 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 climberon tree trunks, up to 20 m above ground. SEASONAL OCCURRENCE: Adults have been found inFebruary – November; copulating pairs in April – May; gravid females in April; teneral adults inApril, June – July, and September. BEHAVIOR: Nocturnal, adults take cover during the dayamong dead leaves and under stones, fallen trees, boards, and in rotten tree-stumps. Adults aregregarious. 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 soft-bodied insect larvae.References: Ciegler (2000), Erwin (1981, 2001), Larochelle & Larivière (2003), Data fromNMNH collection

Carabus (Carabus) granulatus granulatus Linné 1758GRANULATE WORM & SLUG HUNTER

Carabus granulatus Linné, 1758:41Carabus chalybaeus Voet, 1778:70Carabus campestris M.F. Adams, 1817:297Carabus confluens Fischer von Waldheim, 1828:187Carabus parallelus Faldermann, 1835:19Carabus niger Letzner, 1850:73Carabus nigrescens Letzner, 1850:73Carabus rufofemoratus Letzner, 1850:73Carabus virescens Letzner, 1850:73Carabus rubripes Géhin, 1876:15Carabus cupreoaeneus Dalla Torre, 1877:25Carabus nigroaeneus Dalla Torre, 1877:25Carabus rubripes Dalla Torre, 1877:25Carabus viridiaeneus Dalla Torre, 1877:25


Carabus parvicollis Kraatz, 1878:129Carabus forticostis Kraatz, 1878:130Carabus haematomerus Kraatz, 1878:131Carabus fulvipes Géhin, 1885:26Carabus Wimmeli Schulz, 1900:160Carabus expansus Vacher de Lapouge, 1905:304Carabus crimeensis Breuning, 1933:527Carabus granulatulus Mandl, 1955:249Carabus atrocyaneus Lamy, 1973:71Carabus 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 meadowsclose to water in low deciduous forests (often stands of Alnus), swamps, along river banks andlake 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, andSeptember – November. BEHAVIOR: Mostly nocturnal, spring breeder, usually adults take coverduring the day under the loose bark of tree-stumps. Adults are gregarious in winter. Eggs arelaid 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 arepredaceous on snails. Markedly favored by human activities, thus effective as a colonist.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth, et al. (1985), Data fromNMNH collection

Carabus (Carabus) granulatus hibernicus Lindroth 1956CELTIC 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: Onopen moist or wet, clayish ground covered with some vegetation. Along field margins, in cultivatedfields (e.g., hay, potato); at borders of marshes and temporary pools, mud flats, lake shores, andin orchards. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Adults have been found in April – September, andNovember. BEHAVIOR: Nocturnal, adults take cover during the day under stones, pieces ofwood or cardboard, and under dead fallen plants. Markedly favored by human activities, thuseffective as a colonist.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1956, 1961)


Carabus (Tanaocarabus) hendrichsi Bolívar y Pieltain 1967HENDRICHS’ 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 andconiferous forests. MICROHABITAT: On the ground in forests of predominately Pinus hartwegiiLindl, P. strobiformis Engelmann, Abies vejari Marinez, and at the lower parts of the altitudinalrange, 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 walkingor 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 1820MENDERES RIVER WORM & SLUG HUNTER

Carabus maeander Fischer von Waldheim, 1820: pl. 10Carabus incompletus Fischer von Waldheim, 1828:303Carabus Ehrenbergi Fischer von Waldheim, 1829:368Carabus Lapilayi Laporte de Castelnau, 1834:89Carabus hudsonicus Motschulsky, 1866:293Carabus maeander Simoni von Heyden, 1879:163Carabus maeander excatenatus Kraatz, 1880:337Carabus maeander excostatus Kraatz, 1880:338Carabus Lecontei Géhin, 1885:27Carabus obscuratus Géhin, 1885:26Carabus maeander mongolicus Vacher de Lapouge, 1905:306Carabus 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 denselow 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, andirrigation 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; gravidfemales in May; teneral adults in July – August. BEHAVIOR: Diurnal and nocturnal, active in thesunshine among vegetation (especially in spring) or adults take cover on cloudy or cool daysunder 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 theupper zone of brook borders, roadsides, and gravel pits, in vacant lots, on hills, and at the edgeof woods, under well-embedded stones, in the ground at the base of trees, under the loose barkof fallen trees, and under fallen trees. Moderately favored by human activities.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection


Carabus (Archicarabus) nemoralis nemoralis Müller 1764WOODLAND WORM & SLUG HUNTER

Carabus nemoralis O.F. Müller, 1764:21Carabus 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 inhumus 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 vagilitylimited to walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have beenfound in March – December; copulating pairs in April – June; gravid females in May; teneraladults in July – September. BEHAVIOR: Mostly nocturnal, spring breeder, usually adults takecover during the day under stones, boards, stacks of lumber, dead leaves, heaps of hay, andother debris; sometimes active on cloudy days on the ground. Coitus lasts 10 – 54 minutes; eggsare 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 leaflitter, 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 adultsare found swarming on the ground under artificial lights at night. Larval are predaceous onlepidopterous caterpillars. Adults play an important role in the control of slugs, thus they are ofeconomic importance. Markedly favored by human activities (e.g., cultivation and transportationof agricultural and horticultural produce), thus very effective as a colonist.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNHcollection

Carabus (Hemicarabus) serratus Say 1823SERRATE-SHOULDER WORM & SLUG HUNTER

Carabus serratus Say, 1823:77Carabus Lineoatopunctatus Dejean, 1826:77Carabus Tatumi Motschulsky, 1865:293Carabus 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, gravelpits, 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 openforests; in sunny glades in woodlands with fireweed, Epilobium angustifolium L. DISPERSAL ABILITIES:Wing-dimorphic: Macropterous form probably capable of flight. SEASONAL OCCURRENCE: Adults


have been found in March – November, rarely in July; copulating pairs in June; gravid femalesin May; teneral adults in July – September. BEHAVIOR: Mostly nocturnal, sometimes active inthe spring sunshine, usually adults take cover during the day under stones, fallen trees, pieces ofcardboard, 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.). Markedlyfavored by human activities.References: Erwin (1981, 2001), Larochelle (1972, 1975), Larochelle & Larivière (2003), Lindroth(1961), Data from NMNH collection

Carabus (Tanaocarabus) sylvosus Say 1823SILVAN WORM & SLUG HUNTER

Carabus sylvosus Say, 1823:75Carabus 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 shadedmore or less moist ground, in wooded areas such as deciduous (e.g., Quercus), mixed, andconiferous 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 orrunning. SEASONAL OCCURRENCE: Adults have been found in January – December; gravid femalesin 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 foundoverwintering in woodlots under fallen trees and in the ground. Adult are predaceous onlepidopterous pupae and scarabaeids.References: Ciegler (2000), Erwin (1981, 2001), Larochelle & Larivière (2003), Lindroth (1961),Data from NMNH collection

Carabus (Oreocarabus) taedatus agassii LeConte 1850AGASSI’S WORM & SLUG HUNTER

Carabus Agassii LeConte 1850:209Carabus oregonensis LeConte, 1854:16Carabus gladiator Motschulsky, 1866:285Carabus Agassizi von Heyden, 1879:162Carabus taedatus canadicus Roeschke, 1900:70Carabus patulicollis Casey, 1913:57Carabus taedatus montanicus Casey, 1913:58Carabus stocktonensis Casey, 1920:155Carabus 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: MACROHABITAT: Lowlands, mountains, subalpine and alpine zones.MICROHABITAT: On open rather dry, mostly gravelly ground covered with thin low vegetationor dead leaves on moraines, meadows, shrub-steppes, in canyons, on mountain slopes and


cirques, in cultivated fields (e.g., wheat, barley), on sagebrush-steppes, rangelands, in opendeciduous (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. Quickto colonize recently burnt woods. SEASONAL OCCURRENCE: Adults have been found in May –October. BEHAVIOR: Nocturnal, adults take cover during the day under stones and fallentrees. Adults found overwintering in woodlots in rotten tree-stumps. Adult are predaceouson caterpillars. Adults may play an important economic role in the control of the Forest TentCaterpillar (Malacosoma disstria Hubner).References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNHcollectionNotes: Kavanaugh (in litt.), based on a study by one of his former students, informs me thatthe C. taedatus complex breaks out into 4 species. Since this has not yet been published, I reporthere in the sense of Lindroth (1961) followed by Larochelle & Larivière (2003).

Carabus (Oreocarabus) taedatus taedatus Fabricius 1787PINE FOREST WORM & SLUG HUNTER

Carabus taedatus Fabricius, 1787:196Carabus baccivorus Fischer von Waldheim, 1820: pl. 7Carabus seriatus Wiedemann, 1821:109Carabus franciscanus Casey, 1913:58Carabus bicanaliceps Casey, 1920:154Carabus 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; sometimesactive during the day on snow patches.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNHcollectionNotes: 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 herein the sense of Lindroth (1961) followed by Larochelle & Larivière (2003).

Carabus (Aulonocarabus) truncaticollis truncaticollis Eschscholtz 1833BLUNT-COLLARED WORM & SLUG HUNTER

Carabus truncaticollis Eschscholtz, 1833:22Carabus lutschnikianus Basilewsky, 1937:63Carabus 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 thetundra. MICROHABITAT: Open moderately moist, stony or muddy ground covered with meadow-


type vegetation, dwarf birches, or willows. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJune – August; larvae in July – August. BEHAVIOR: Larvae dwell under moss carpets. Adults arepredaceous on dipterous larvae.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNHcollection

Carabus (Megodontus) vietinghoffi vietinghoffi Adams 1812VIETINGHOFF’S WORM & SLUG HUNTER

Carabus Vietinghoffi Adams, 1812:170Carabus Stschegolewi Poppius, 1906:17Carabus alaskanus Obydov, 1996:85

Distribution: Native, New World. CANADA – NT, YT; USA – AK.Way of Life: MACROHABITAT: Lowlands, 12 – 500 meters altitude. MICROHABITAT: On shadedor 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) 1801BOUND WORM & SLUG HUNTER

Tachypus vinctus Weber, 1801:42Carabus interruptus Say, 1823:76Carabus ligatus Germar, 1824:6Carabus Carinatus Dejean, 1826:80Carabus 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 shadedor half-shaded wet, clayish or muddy ground covered with dead leaves close to water in flood-plain forests (e.g., Liriodendron, Liquidambar), low deciduous forests, and swamp-forests. DISPERSAL


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; inrotten tree-stumps and under the loose bark of fallen trees. Adults are able to survive winterinundation of their hibernation site. Adult are scavengers on dead fish, meat, and fowl. Adultsoccasionally found walking in to lights at night.References: Ciegler (2000), Erwin (1981, 2001), Larochelle & Larivière (2003), Data fromNMNH collection


Ceroglossina Lapouge 1927NEAUSTRAL 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 notbased on breeding potentials, rather color and slight morphological variation, as well asimperfectly known ranges. A thorough genomics study would greatly benefit the taxonomy ofthis group.Adelphotaxon: Carabina Latreille 1802.Geographic status: Neaustral.References: Erwin (2001), Jiroux (1996), Lorenz (2005)

Ceroglossus Solier 1848NEAUSTRAL MAGNIFICENT BEETLES

(Plate 18)

Ceroglossus Solier, 1848:59Ceroglossa 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 notbased on breeding potentials. Adelphotaxon: Calosoma Weber, 1801Type species: Carabus suturalis Fabricius, 1775Distribution: Argentina, Chile.Habitat: Adults of this genus are found exclusively in forested or woodlot areas, even smallcoppice, principally in Nothofagus and Araucaria forests, woodlots, their edges, or in thickets withproximity to farms and ranches. They are dwellers of the ground, hiding themselves during theday 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 ofinteresting color forms throughout it range. It is ripe for serious molecular studies that will helpus understand breeding potentials and lead to a stabilization of the species taxonomy. Thetreatment below does not contain the most recent barrage of subspecies descriptions publishedin obscure journals; they will be treated in the first supplementum.

Ceroglossus buqueti andestus Kraatz-Koschlau 1887RAMPART MAGNIFICENT BEETLE

Ceroglossus Buqueti andestus Kraatz-Koschlau, 1887:193Carabus Cyaneus Faz, 1925:22Carabus Salbachi Faz, 1925:23

Distribution: Native, New World. CHILE.


Way of Life: MACROHABITAT: Lowlands to midlands, 500 – 1100 meters altitude, in forests andgroves 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 fallentree; 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 1996ARGENTINE 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 andwoodlots 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: Seedescription under genus, above.References: Erwin (2001), Jiroux (1996), Data from NMNH collection

Ceroglossus buqueti arieli Jiroux 1996ARIEL’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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJanuary. BEHAVIOR: See description under genus, above. Cohabits with adults of C. magellanicusmagellanicus.References: Erwin (2001), Jiroux (1996)

Ceroglossus buqueti arriagadai Jiroux 1996ARRIAGADA’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 towalking 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)


Ceroglossus buqueti breuningi Heinz & Jiroux 2001BURENING’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 Araucariaforests 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) 1835BUQUET’S MAGNIFICENT BEETLE

Carabus Buqueti Laporte de Castelnau, 1835:158Carabus chilensis Guérin-Méneville, 1835: pl. 3Carabus Valdiviae Hope, 1837:128Carabus Valdiviae Gerstaecker, 1858:431Ceroglossus Valdivianus Morawitz, 1886:439Ceroglossus 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 thicketsand coppice of Nothofagus, Gevuina, and Persea spp. MICROHABITAT: Ground-dwelling. DISPERSAL


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) 1838CHILOÉ ISLAND MAGNIFICENT BEETLE

Carabus chiloensis Hope, 1838:128Carabus insularis Hope, 1838:129Ceroglossus dorsiger Motschulsky, 1865:284Carabus elegantissimus Reed, 1874:53Ceroglossus Buqueti chloridus Géhin, 1885:55Ceroglossus Buqueti Hopei Kraatz-Koschlau, 1885:420Ceroglossus Buqueti castroensis Kraatz-Koschlau, 1886:419Ceroglossus Monttianus Morawitz, 1886:437Ceroglossus Buqueti punctulatus Morawitz, 1886:438Carabus Porteri Faz, 1925:14Carabus Buqueti Claudii Faz 1925:22Ceroglossus buqueti penai Van Dorsselaer, 1955:88


Distribution: Native, New World. ARGENTINA – CHUBUT, NEUQUÉN; CHILE – ISLA CHILOE.Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1219 meters altitude, in forestwoodlots across a broad territory, in biotopes of very different kinds, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Adults have been found in January – February, andDecember. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996), Data from NMNH collection

Ceroglossus buqueti cupreus (Faz) 1925COPPERY MAGNIFICENT BEETLE

Carabus Sybarita Cupreus Faz, 1925:23Ceroglossus psittacus Gerstaecker, 1858:425Ceroglossus sybarita calvus Géhin, 1885: pl. 54Ceroglossus sybarita tomentosus Kraat-Koschlau, 1885:421

Distribution: Native, New World. CHILE – ISLA CHILOE.Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Ground-dwelling. DISPERSAL


OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996, 2001)

Ceroglossus buqueti deuvei Jiroux 1996DEUVE’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 undergenus, above. Cohabits with C. chilensis ficheti and C. magellanicus dolhemi.References: Erwin (2001), Jiroux (1996), Data from NMNH collection

Ceroglossus buqueti lepidus Kraatz-Koschlau 1891YELLOWISH MAGNIFICENT BEETLE

Ceroglossus Buqueti lepidus Kraatz-Koschlau, 1891:11Carabus Similis Faz, 1925:22Carabus Similis Elegans Faz, 1925:22Carabus Similis Incertus Faz, 1925:22Ceroglossus 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 SouthernBeech (Nothofagus), MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence


flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound 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 1998LORENZ’S MAGNIFICENT BEETLE

Carabus curtus Germain, 1895:58Ceroglossus buqueti lorenzi Jiroux, 1998:171

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Ground-dwelling. DISPERSAL


OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1998)

Ceroglossus buqueti magdalenaensis Jiroux 1996MAGDALENA 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 wetisland forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January and March. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996)

Ceroglossus buqueti peladosus Kraatz-Koschlau 1887PELADA MOUNTAINS MAGNIFICENT BEETLE

Ceroglossus Valdiviae peladosus Kraatz-Koschlau, 1887:358Ceroglossus Buqueti inexpectatus Kraatz-Koschlau, 1890:252Carabus Similis Politus Faz, 1925:22

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


Ceroglossus buqueti subnitens Kraatz-Koschlau 1885SLIGHTLY SHINY MAGNIFICENT BEETLE

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

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Lowlands to midlands, 74 – 1000 meters altitude, in thickets ofSouthern 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 1858LUXURIOUS 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, mixedforests. MICROHABITAT: Ground-dwelling at borders of lakes. DISPERSAL 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 1888ANGOL MAGNIFICENT BEETLE

Ceroglossus gloriosus chilenicus Kraatz-Koschlau, 1888:109Ceroglossus gloriosus angolicus Kraatz-Koschlau 1888:110Carabus Gloriosus percuprinus Faz, 1925:19Carabus Gloriosus Subviridus Faz, 1925:19Carabus Gloriosus Jaffueli Faz, 1925:19Carabus Gloriosus Viridis Faz, 1925:19Carabus 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 towalking 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


Ceroglossus chilensis chilensis (Eschscholtz) 1829CHILEAN MAGNIFICENT BEETLE

Carabus chilensis Eschscholtz, 1829:9Ceroglossus ignitus Géhin, 1885:97Ceroglossus Lossbergi Kraatz-Koschlau, 1887:195Ceroglossus 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 Bio-Bio, about thirty kilometres from the ocean. The topography of the region at Hualqui on theBio Bio River varies, starting from the west with coastal lowland marshes, then crossing a lowcoastal mountain (dominated by Araucaria pines) range running north to south, followed by arich fertile interior valley, and finally the Andean mountain range in the east. MICROHABITAT:Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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) 1875COLCHAGUA MAGNIFICENT BEETLE

Carabus colchaguensis Reed, 1875:225

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Midlands, 800 meters altitude, in Southern Beech forests [Nothofagusalpina 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 1887BLUE-COLLARED MAGNIFICENT BEETLE

Ceroglossus cyanicollis Kraatz, 1887:230Carabus Chilensis Germanus Faz, 1925:18

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


Ceroglossus chilensis evenoui Jiroux 1996EVENOU’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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary. BEHAVIOR: See description under genus, above. Cohabits with members of C. buquetisubnitens in the southern end of its range.References: Erwin (2001), Jiroux (1996)

Ceroglossus chilensis fallaciosus Kraatz 1880DECEPTIVE MAGNIFICENT BEETLE

Ceroglossus chilensis fallaciosus Kraatz, 1880:56Ceroglossus pyrilampes Morawitz, 1886:406Ceroglossus gloriosus incertus Kraatz-Koschlau, 1888:312

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in mixed forests, small isolatedthickets 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 collectionNotes: Found in between Vichuquen in the north and Arauco in the south. This area is heavilypolluted and undergoing deforestation; conifers are replacing the Beech forests.

Ceroglossus chilensis ficheti Jiroux 1996FICHET’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 ofNothofagus 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


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)


Ceroglossus chilensis germaini Jiroux 1996GERMAIN’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 arucariaforests 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 undergenus, above. Cohabits with C. buqueti chiloensis and C. darwini reedi.References: Erwin (2001), Jiroux (1996)

Ceroglossus chilensis gloriosus Gerstaecker 1858GLORIOUS MAGNIFICENT BEETLE

Ceroglossus gloriosus Gerstaecker, 1858:429Ceroglossus gloriosus carinulatus Motschulsky, 1865:284Ceroglossus gloriosus Pradieri Géhin, 1885:55Ceroglossus gloriosus unicostulatus Géhin, 1885:55Ceroglossus gloriosus dorsosolutus Morawitz, 1886:403Ceroglossus gloriosus temucensis Kraatz-Koschlau, 1890:257Carabus Aeneovirescens Aeneus Faz, 1925:20Carabus Aeneovirescens Faz, 1925:20Ceroglossus chilensis gloriosus mallecoanus Vacher de Lapouge, 1928:7Carabus diversicostatus Dallas, 1828:81Ceroglossus chilensis Porteri Dallas, 1929:351Ceroglossus debilicostis Dallas, 1936:74

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Lowlands to midlands, 4 – 933 meters altitude, in thickets, coppiceand groves. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound 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 1886KRAATZ’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 coppiceof 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


adults are common and cohabit with C. buqueti buqueti and C. magellanicus candens, which also areextremely well represented.References: Erwin (2001), Jiroux (1996)

Ceroglossus chilensis latemarginatus Kraatz-Koschlau 1889TILED-MARGIN MAGNIFICENT BEETLE

Ceroglossus gloriosus latemarginatus Kraatz-Koschlau, 1889:110Ceroglossus dynastes Born, 1898:656Ceroglossus 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 coniferousforests. MICROHABITAT: Ground-dwelling on forested slopes. DISPERSAL ABILITIES:Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL

OCCURRENCE: Adults have been found in February and December. BEHAVIOR: See descriptionunder genus, above.References: Erwin (2001), Jiroux (1996), Data from NMNH collection

Ceroglossus chilensis legrandi Heinz & Jiroux 2001LEGRAND’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 dryNothofagus 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 2001SOUTHERN 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 Nothofagusforests. MICROHABITAT: Ground dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJanuary. BEHAVIOR: See description under genus, above. Cohabits with C. darwini patagonensis.References: Erwin (2001), Heinz & Jiroux (2001)


Ceroglossus chilensis mochae (Reed) 1874MOCHA ISLAND MAGNIFICENT BEETLE

Carabus Mochae Reed, 1874:54Carabus mochae conchyliatus Géhin, 1875:56Carabus 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 - MochaIsland 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: Seedescription under genus, above. This is the only form of Ceroglossus to inhabit this isolated partof the Cordillera de Nahuelbuta.References: Erwin (2001), Jiroux (1996), Data from NMNH collection

Ceroglossus chilensis nigritulus Mandl 1977MANDL’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. nigrituluspredominate near Lake Menendez, whereas members of C. d. minutissimus are more frequent inthe remainder of the park.References: Erwin (2001), Jiroux (1996)

Ceroglossus chilensis pseudopatagoniensis Heinz & Jiroux 2001FALSE 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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJanuary – February. BEHAVIOR: See description under genus, above. Cohabits with C. buquetichiloensis and C. darwini patagoniensis.References: Erwin (2001), Heinz & Jiroux (2001)


Ceroglossus chilensis seladonicus Kraatz-Koschlau 1887MOON MAGNIFICENT BEETLE

Ceroglossus gloriosus seladonicus Kraatz-Koschlau, 1887:194Ceroglossus gloriosus hypocrita Kraatz-Koschlau, 1887:194Carabus Aeneovirescens Ruizi Faz, 1925:20Carabus Aeneovirescens Plagiarus Faz, 1925:20Ceroglossus 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 coppiceof Nothofagus alpina Poepp. & Endl., N. pumilio (Poepp. et Endl.) Krasser, and N. dombeyi(Mirb.) Blume, etc. MICROHABITAT: In the region of Caburga, Cordillera de LonquimayaNeltume, this subspecies is found in large groves of these southern beeches under debrison the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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 1900SOLIER’S MAGNIFICENT BEETLE

Ceroglossus Solieri Roeschke, 1900:61Ceroglossus 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, andimpenetrable 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 – February, October, and December. BEHAVIOR:See description under genus, above. Cohabits with members of C. buqueti chiloensis, C. suturalispuxel 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 1885VILLARICA MAGNIFICENT BEETLE

Ceroglossus gloriosus villaricensis Kraatz-Koschlau, 1885:431Carabus Gloriosus Villaricensis Capuccinus Faz, 1925:19Carabus Gloriosus Darwinii Reedjuniori Faz, 1925:19

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Lowlands, 135 – 400 meters altitude, in a cordillera with difficultaccess, 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


under genus, above. Adults are rather common here often seen crossing the road. Adults takecover under debris in forested thickets.References: Erwin (2001), Jiroux (1996), Data from NMNH collection

Ceroglossus darwini darwini (Hope) 1837DARWIN’S MAGNIFICENT BEETLE

Carabus Darwini Hope, 1837:129Ceroglossus indiconotus Solier, 1849:127Ceroglossus melanopterus Gerstaecker, 1858:429Ceroglossus Darwini bimarginatus Kraatz-Koschlau, 1886:418Ceroglossus indiconotus disputatus Kraatz-Koschlau, 1887:228Ceroglossus indiconotus opacus Kraatz-Koschlau, 1887:228Ceroglossus Darwini chonchicus Kraatz-Koschlau, 1887:356Ceroglossus darwini auroviridis Bruening, 1943:111Ceroglossus 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in July. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996)

Ceroglossus darwini jirouxi Deuve 1995JIROUX’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 RioManihuales. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996)

Ceroglossus darwini mequignoni Heinz & Jiroux 2001MEQUIGNON’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 towalking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR:See description under genus, above.References: Erwin (2001), Heinz & Jiroux (2001)


Ceroglossus darwini patagoniensis Breuning 1931PATAGONIAN 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 theValle de Lago Blanco was completey burned in 1940 to raise sheep. All that is left now aregrasslands 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 1886REED’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 arucariaforests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Jiroux (1996)

Ceroglossus guerini Géhin 1885GUERIN’S MAGNIFICENT BEETLE

Ceroglossus Guerini Géhin, 1885:55Ceroglossus explanatus Breuning, 1943:24Ceroglossus intermedius Breuning, 1943:25

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 859 meters altitude in thicketsand coppice of Nothofagus, Gevuina, and Persea spp. MICROHABITAT: Ground-dwelling with denselyshaded understory. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description undergenus, above. Cohabits with C. buqueti buqueti.References: Erwin (2001), Jiroux (1996)

Ceroglossus magellanicus araucanus Kraatz-Koschlau 1886AURACANA 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


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 1996BOEUF’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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January. BEHAVIOR: See description under genus, above. Cohabits with C. chilensisgloriosus at Victoria and Tolhuaca, and with C. buqueti lepidus at Tolhuaca.References: Erwin (2001), Jiroux (1996)

Ceroglossus magellanicus bouchardi Jiroux 1996BOUCHARD’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 towalking 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 1989CATABURGA 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)


Ceroglossus magellanicus candens Géhin 1885BRIGHT MAGNIFICENT BEETLE

Ceroglossus candens Géhin, 1885:96Ceroglossus bicostulatus Kraatz-Koschlau, 1885:428Ceroglossus Morawitzi Kraatz, 1887:236

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Lowlands, sea level – 43 meters altitude, in thickets and coppiceof 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 1998DAVIDSON’S MAGNIFICENT BEETLE

Ceroglossus magellanicus davidsoni Jiroux, 1998:170

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Ground-dwelling. DISPERSAL


OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. Probably cohabitswith C. buqueti peladosus in the forests of the Cordillera Pelada.References: Erwin (2001), Jiroux (1998)

Ceroglossus magellanicus dolhemi Jiroux 1996DOLHEM’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: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: Seedescription under genus, above. Cohabits with C. buqueti deuvei and C. chilensis ficheti.References: Erwin (2001), Jiroux (1996)

Ceroglossus magellanicus gerstaeckeri Morawitz 1886GERSTAECKER’S MAGNIFICENT BEETLE

Ceroglossus Gerstaeckeri Morawitz, 1886:429Ceroglossus tenebriculus Kraatz-Koschlau, 1887:357 (?)Carabus Magnus Faz, 1925:11Carabus magnus vicinus Faz, 1925:11


Carabus magnus caputolivae Faz, 1925:11Carabus magnus imitator Faz, 1925:11Ceroglossus 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: Seedescription under genus, above.References: Erwin (2001), Jiroux (1996), Data from NMNH collection

Ceroglossus magellanicus magellanicus Géhin 1885SOUTHERN-MOST MAGNIFICENT BEETLE

Ceroglossus magellanicus Géhin, 1885:56Carabus valdiviae aerosus Faz, 1925:10Carabus valdiviae proximus Faz, 1925:10Carabus magnus fusco Faz, 1925:11

Distribution: Native, New World. CHILE.Way of Life: MACROHABITAT: Lowlands, sea level – 211 meters altitude, in thickets, coppiceand groves. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in October. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996)

Ceroglossus magellanicus newtoni Jiroux 1999NEWTON’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 descriptionunder genus, above.References: Erwin (2001), Jiroux (1999)

Ceroglossus magellanicus ortrudheinzae Heinz & Jiroux 2001ORTRUDHEINZ’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


vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJanuary and December. BEHAVIOR: See description under genus, above. Cohabits with C. chilensisgloriosus and C. buqueti buqueti.References: Erwin (2001), Heinz & Jiroux (2001)

Ceroglossus magellanicus similis Kraatz-Koschlau 1885FAZ’S MAGNIFICENT BEETLE

Ceroglossus similis Kraatz-Koschlau, 1885:435Carabus Hyocrita Celadonicus galbinus Faz, 1925:10Carabus Hyocrita Celadonicus aestectus Faz, 1925:10Carabus 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) 1895OCHSEN’S MAGNIFICENT BEETLE

Carabus Ochseni Germain, 1895:40Ceroglossus 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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996)

Ceroglossus speciosus Gerstaecker 1858SHOWY MAGNIFICENT BEETLE

Ceroglossus speciosus Gerstaecker, 1858:438Carabus Speciosus Oyarzuni Faz, 1925:14

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


Ceroglossus suturalis ancudanus Morawitz 1886ANCUD MAGNIFICENT BEETLE

Ceroglossus suturalis ancudanus Morawitz, 1886:387Ceroglossus 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: Adultshave 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 1886OLIVACEOUS MAGNIFICENT BEETLE

Ceroglossus suturalis olivaceus Kraatz-Koschlau, 1886:420Ceroglossus suturalis suturelevatus Kraatz-Koschlau, 1888:311Ceroglossus 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 impenetrablemixed 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 1996PUXEL 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 Alerceforests [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 averagetemperatures 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January. BEHAVIOR: See description under genus, above.References: Erwin (2001), Jiroux (1996), www.conifers.org


Ceroglossus suturalis suturalis (Fabricius) 1775SEAMED MAGNIFICENT BEETLE

Carabus suturalis Fabricius, 1775:238Carabus Reichei Guérin-Méneville, 1839:297Carabus Dubitabilis Faz, 1925:14Carabus Suturalis Magallanicus Faz, 1925:15Carabus 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 nearthe sea. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. 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 1834SNAIL-EATING BEETLES

Cychrini Laporte de Castelnau, 1834:86

Number of worldwide genera: Six.Number of Western Hemisphere genera: Three.Taxonomy: Stable. Adelphotaxon: Carabini Latreille 1802Geographic 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 epipleuralsurfaces against the lateral margin of the tergites (Roeschke, 1907).References: Erwin (2001), Lorenz (2005), (Roeschke, 1907)

Cychrus Fabricius 1794RARE SNAIL-EATING BEETLES

(Plate 19)

Cychrus Fabricius, 1794:440Cychrys Fabricius, 1794:440Kryptocychrus Cavazzuti, 1997:75See 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 notbased on breeding potentials. Likely, there are many new species to be discovered in China.Adelphotaxon: Cychropsis Boileau 1901


Type species: Carabus rostratus Fabricius, 1775Distribution: Holarctic: Nearctic (two species), Palearctic (79 species).Habitat: According to Lindroth (1961), most species of this genus are confined to forestedcountry. 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 1878HEMPHILL’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 orbrushy forest fragments (e.g. Alnus, Philadelphicus) growing along rivers in canyons. Also, foundin mixed coniferous forest with Populus and Abies. MICROHABITAT: On shaded rather dry groundwith dense vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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 collectionsNotes: Both forms, C. h. hemphillii and C. h. rickseckeri have representative specimens labeled LoganPass, Utah. This area ranges from 1357 – 3050 meters altitude. Perhaps the two subspecies arealtitudinally separated where their ranges overlap.

Cychrus hemphillii rickseckeri LeConte 1884RICKSECKER’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 andconiferous forests often along streams. MICROHABITAT: On shaded rather dry ground withsome vegetation DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 orlarge fallen branches, or trunks on the ground. Larvae hibernate.References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS,NMNH collectionsNotes: See note above.

Cychrus tuberculatus T.W. Harris 1839TUBERCULATE RARE SNAIL-EATING BEETLE

Cychrus tuberculatus T.W. Harris, 1839:200Cychrus pustulosus Casey, 1905:160

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


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 limitedto 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 arepredaceous on snails. Larvae hibernate.References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNHcollection

Scaphinotus Dejean 1826SNAIL-EATING BEETLES

(Plate 20)

Scaphinotus Dejean, 1826:17Scaphinopterus Fischer von Waldheim, 1828:136Irichroa Newman, 1838:385Scaphonotus Agassiz, 1847:960Analobus Gistel, 1857:90Megalridia Casey, 1920:175Nomaretus LeConte, 1853:399 [valid subgenus for native Nearctic species]Stenocantharis Gistel, 1857:92 [valid subgenus for native Nearctic species]Phemphus Motschulsky, 1866:312Brennus 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 1826Type species: Carabus elevatus Fabricius, 1787Distribution: Nearctic.Habitat: According to Lindroth (1961), most species of this genus are confined to forestedcountry, many to mountainous areas. Adults are nocturnal; most species hibernate as larvae, atleast in part. Adults are adapted to eating land snails, although Noonan (1967) observed adultsof 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 withbetter distributional data, as well as their population genetics.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Noonan (1967)

Scaphinotus (Steniridia) aeneicollis (Beutenmüller) 1903COPPERY-COLLARED SNAIL-EATING BEETLE

Cychrus aeneicollis Beutenmüller, 1903:515Cychrus purpuratus Beutenmüller, 1918:89


Distribution: Native, New World. USA – NC.Way of Life: MACROHABITAT: Midlands to uplands, 914 – 1980 meters altitude, in coniferousforests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: In southern Appalachiaalong streams in ravines on shaded moist ground. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in May – October. BEHAVIOR: Nocturnal, taking cover in the day under loose tree bark.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNHcollection

Scaphinotus (Steniridia) andrewsi amplicollis (Casey) 1920WIDE-COLLARED SNAIL-EATING BEETLE

Irichroa violacea amplicollis Casey, 1920:174Irichroa andrewsi reflexa Casey, 1924:22

Distribution: Native, New World. USA – NC, VA.Way of Life: MACROHABITAT: Midlands to uplands, 1220 – 1524 meters altitude, in coniferousforests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moderatelymoist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 fromNMNH collection

Scaphinotus (Steniridia) andrewsi andrewsi (T.W. Harris) 1839ANDREWS’ 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 mixedforests (Quercus, Pinus, Cornus, and Carya spp.). MICROHABITAT: On shaded moist ground alongthe banks of woodland streams on the southern Appalachia piedmont. A specimen collected inVirginia in 1943 is labeled “Oak-Chestnut.” DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJune – August, October and December. BEHAVIOR: Nocturnal, taking cover in the day underloose tree bark, in rotten wood, or in tree hollows; adults are gregarious in autumn. Adults eatberries and snails. Regular climber on tree trunks while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data fromNMNH collection

Scaphinotus (Steniridia) andrewsi barksdalei (Valentine) 1936BARKSDALE’S SNAIL-EATING BEETLE

Steniridia andrewsi barksdalei Valentine, 1936:230

Distribution: Native, New World. USA – NC.


Way of Life: MACROHABITAT: Midlands to uplands, 914 – 1676 meters altitude, in coniferousforests (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 descriptionunder genus, above.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1936), Data from NMNHcollection

Scaphinotus (Steniridia) andrewsi darlingtoni (Valentine) 1935DARLINGTON’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 coniferousforests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moist groundon ridges and in ravines. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagilitylimited 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 fromNMNH collection

Scaphinotus (Steniridia) andrewsi germari (Chaudoir) 1861GERMAR’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 CumberlandMountains, in mixed forests (e.g., Quercus spp., Acer spicatum Lam., Osytra spp.) on slopes and in ravineswith a dense growth of eastern hemlock (Tsuga canadensis (L.) Carr.), rhododendron (R. catawbienseBoursault and R. maximum L.) and mountain laurel (Kalmia latifolia L.). MICROHABITAT: On shadedmoderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 arepredators on snails. Regular climber on fallen tree trunks and cave walls while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data fromNMNH, UASM, collections

Scaphinotus (Steniridia) andrewsi montanus (Valentine) 1935MOUNTAIN 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


eastern hemlock (Tsuga canadensis (L.) Carr.), rhododendron (R. catawbiense Boursault and R. maximumL.) and mountain laurel (Kalmia latifolia L.). MICROHABITAT: On shaded moist somewhat rockyground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. 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 fromNMNH collection

Scaphinotus (Steniridia) andrewsi mutabilis (Casey) 1920CHANGELING SNAIL-EATING BEETLE

Irichroa mutabilis Casey, 1920:173Irichroa mutabilis longicollis Casey, 1920:173Irichroa 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., Quercusspp., 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 walkingor 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 NMNHcollection

Scaphinotus (Steniridia) andrewsi nantahalae (Valentine) 1936NANTAHALA 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: Adultshave been found in April – October. BEHAVIOR: Nocturnal, taking cover in the day in rottentree stumps and fallen trees, occasionally under moss carpets. Frequent climber on tree stumpsand fallen trees while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1936), Data from NMNH,UASM collections

Scaphinotus (Steniridia) andrewsi parvitarsalis (Valentine) 1935SMALL-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


thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJune – October. BEHAVIOR: Nocturnal, taking cover in the day under loose bark of tree stumpsand fallen trees. Frequent climber on tree stumps and fallen trees while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data fromNMNH collection

Scaphinotus (Steniridia) andrewsi saludae (Valentine) 1936HEALTHY 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 undergenus, above.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1936), Data from NMNHcollection

Scaphinotus (Steniridia) andrewsi waldensius (Valentine) 1935WALDEN 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 mixedforest. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inOctober. BEHAVIOR: See description under genus, above.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data fromNMNH collection

Scaphinotus (Neocychrus) angulatus (T.W. Harris) 1839ANGULATE SNAIL-EATING BEETLE

Cychrus angulatus T.W. Harris, 1839:200Scaphinotus 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, intemperate coniferous rainforest (Pseudosuga). MICROHABITAT: On shaded moderately moistground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. Occasionally climbing on tree trunks. SEASONAL OCCURRENCE: Adults have been foundin April – December. BEHAVIOR: Nocturnal, taking cover in the day under old fallen trees andin 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


Scaphinotus (Stenocantharis) angusticollis (Fischer von Waldheim) 1823NARROW-COLLARED SNAIL-EATING BEETLE

Cychrus angusticollis Fischer von Waldheim, 1823: pl. 46Pemphus angusticollis nigripennis Roeschke, 1907:167Scaphinotus 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 metersaltitude, in forests (coniferous, mixed, and deciduous), and cultivated fields MICROHABITAT:Shaded moist ground often along streams. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inMarch – 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 tree-stumps, 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 onshrubs, tree-stumps, and fallen trees up to 20 m above ground while foraging. Adults have beenfound over wintering in old tree-stumps and fallen trees. Sometimes found in and aroundhuman settlements.References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH,UASM collections

Scaphinotus (Neocychrus) behrensi (Roeschke) 1907BEHRENS’ SNAIL-EATING BEETLE

Neocychrus Behrensi Roeschke, 1907:199Scaphinotus 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 temperateconiferous rainforest (Pseudosuga). MICROHABITAT: On shaded moderately moist ground. DISPERSAL

ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Occasionallyclimbing on tree trunks. SEASONAL OCCURRENCE: Adults have been found in May – July, andSeptember. 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) 1823BILOBED 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


roadsides. MICROHABITAT: Shaded or open moist or wet ground covered with deep leaf litter ormoss, often along rivers, lakes, and the seashore, and in gravel pits surrounded by spruce forests aswell as in Sphagnum-bogs and fens. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility 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 theday under dead leaves, fallen trees, stones, or the loose bark of fallen trees. Frequent climber andgood burrower while foraging. Larval habitat: in leaf litter. Adult are predaceous on snails.References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNHcollection

Scaphinotus (Brennus) bullatus Van Dyke 1924INFLATED SNAIL-EATING BEETLE

Scaphinotus subtilis bullatus Van Dyke, 1924:3Scaphinotus subtilis grandis Van Dyke, 1924:4

Distribution: Native, New World. USA – CA.Way of Life: MACROHABITAT: Lowlands to mountains, 304 – 2286 meters altitude, in coniferousand mixed forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January – March, June – July, and September – November. BEHAVIOR: See descriptionunder genus, above.References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Van Dyke, (1924),Data from CAS, NMNH collections

Scaphinotus (Nomaretus) cavicollis (LeConte) 1859CONCAVE-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


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) 1853HEART-SHAPED THORAX SNAIL-EATING BEETLE

Cychrus cordatus LeConte, 1853:399Brennus cordatus vernicatus Casey, 1920:183Brennus cordatus rufitaris Casey, 1920:184

Distribution: Native, New World. USA – CA.


Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 800 meters altitude, in coastalmixed forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January – March, May – September, and November – December. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS,NMNH collections

Scaphinotus (Brennus) crenatus (Motschulsky) 1859NOTCHED SNAIL-EATING BEETLE

Cychrus crenatus Motschulsky, 1859:161Cychrus striatus LeConte, 1859:69Brennus gentilis Casey, 1897:322Brennus productus Casey, 1914:29Brennus 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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJanuary – October, and December. BEHAVIOR: Nocturnal, taking cover in the day under stonesand fallen trees.References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS,NMNH, UASM collections

Scaphinotus (Brennus) cristatus (T.W. Harris) 1839RIDGED SNAIL-EATING BEETLE

Cychrus cristatus T.W. Harris, 1839:200Cychrus reticulates Motschulsky, 1850:90Brennus basalis Casey, 1897:311Brennus duplicatus Casey, 1897:312

Distribution: Native, New World. USA – CA, OR.Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1150 meters altitude, in mixedand 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


Scaphinotus (Nomaretus) debilis alpinus (Beutenmüller) 1903ALPINE 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. Occasionallyclimbing on tree trunks. SEASONAL OCCURRENCE: Adults have been found in May – June, andAugust – September. BEHAVIOR: Nocturnal, taking cover in the day under stones, loose bark oftrees, and in leaf litter.References: Erwin (2001), Darlington (1931), Larochelle & Larivière (2003), Data from NMNHcollection

Scaphinotus (Nomaretus) debilis debilis (LeConte) 1853WEAKLY 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., Picearubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moderately moist ground rich inleaf litter and humus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 oftrees. Occasional climber while foraging. Larvae found in deep leaf litter. Larvae hibernate.References: Erwin (2001), Darlington (1931), Larochelle & Larivière (2003), Data from NMNHcollection

Scaphinotus (Scaphinotus) elevatus coloradensis Van Dyke 1907COLORADO 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 moderatelymoist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor 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 collectionNotes: See below under the nominate subspecies.


Scaphinotus (Scaphinotus) elevatus elevatus (Fabricius) 1787ELEVATED 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, onmoderately 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 theday under stones, fallen tree-stumps, fallen trees, the loose bark of trees and tree-stumps, deadleaves, 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 collectionNotes: 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 forgenomics. 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 1844FIERY SNAIL-EATING BEETLE

Scaphinotus flammeus Haldeman, 1844:54Cychrus 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 smallpond in Kansas, in open range country. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inApril – November, teneral adults in May – June. Larvae hibernate. BEHAVIOR: Nocturnal, takingcover in the day in leaf litter and under driftwood.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollectionNotes: See above under the nominate subspecies.

Scaphinotus (Scaphinotus) elevatus lengi Van Dyke 1938LENG’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 southwesternVirginia. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless


thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJuly. 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 NMNHcollectionNotes: See above under the nominate subspecies.

Scaphinotus (Scaphinotus) elevatus neomexicanus Van Dyke 1924NEW 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 thusvagility 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 NMNHcollection

Scaphinotus (Scaphinotus) elevatus tenebricosus Roeschke 1907PERFORATE 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: Onshaded wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto walking or running. SEASONAL OCCURRENCE: Adults have been found in May – June, andOctober, teneral adults in May. BEHAVIOR: Nocturnal, taking cover in the day under loose barkof fallen trees. Occasional climber while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollectionNotes: See above under the nominate subspecies.

Scaphinotus (Nomaretus) fissicollis (LeConte) 1853CLEFT-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


OCCURRENCE: Adults have been found in March – June, and August – November, teneral adultsin April and May. BEHAVIOR: See description under genus, above.References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH,UASM collections


Scaphinotus (Steniridia) guyotii (LeConte) 1866GUYOT’S SNAIL-EATING BEETLE

Cychrus Guyotii LeConte, 1866:363Cychrus Angelli Beutenmüller, 1918:89Scaphinotus 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 higheraltitudes in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir), at lower altitudesin mixed and deciduous forests (e.g., Quercus spp., Acer spicatum Lam.), on slopes and in ravineswith a dense growth of eastern hemlock (Tsuga canadensis (L.) Carr.), rhododendron (R. catawbienseBoursault and R. maximum L.) and mountain laurel (Kalmia latifolia L.). MICROHABITAT: On cool,shaded moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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 arepredaceous on snails. Occasional climber while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNHcollection

Scaphinotus (Stenocantharis) hatchi Beer 1971HATCH’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, (Tsugamertensiana (Bong.) Sarg.). MICROHABITAT: On shaded moist ground with sparse vegetation (e.g. Vacciniumscoparium Leiberg), most abundant in broad depressions of the region with more moisture. DISPERSAL


OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Nocturnal, taking cover in theday 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), Datafrom CAS, NMNH, UASM collections

Scaphinotus (Maronetus) hubbardi (Schwarz) 1895HUBBARD’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 deepleaf litter and branches. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto 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 underloose bark of fallen trees. Occasional climber while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections


Scaphinotus (Maronetus) imperfectus (G. Horn) 1860IMPERFECT 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 coniferousforests, and in thickets (e.g. Rhododendron) along streams. MICROHABITAT: On shaded moistground with rich leaf litter and humus. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inMay – November. BEHAVIOR: Nocturnal, taking cover in the day in leaf litter, under stones, andunder loose bark of fallen trees. Occasional climber while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

Scaphinotus (Maronetus) incompletus (Schwarz) 1895INCOMPLETE 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 limitedto 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 1988FORGOTTEN 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: Ground-dwelling in moist, well shaded areas. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility 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 medium-sized 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 inNewton 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 dramaticbluffs and steep hillsides. Geology is a mix of limestone/ dolomite with prominent sandstonecliffs and escarpments. Forest covers are predominantly mature deciduous, with mixed shortleafpine on high points. Land use: pasture/chicken farms/protected forests in the National Riverand Ozark National Forests, which is where the specimen localities are located. Lost Valley isrich beech maple woodland in a valley with a cave at one end. Alum Cove and the area south ofMt. 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 bridgethat I suspect is where that paratype came from because that is where everybody goes. Apparently


Scaphinotus infletus is sympatric with Scaphinotus cavicollis and S. fissicollis, though I don’t see anyindication in our collection that any of them were collected at the same localities.”

Scaphinotus (Brennus) interruptus (Ménétriés) 1843MÉNÉTRIÉS’ SNAIL-EATING BEETLE

Cychrus interruptus Ménétriés, 1843:54Cychrus constrictus LeConte, 1853:398Cychrus dissolutus Schaum, 1863:72Brennus politus Casey, 1897:330Brennus sinuatus Casey, 1897:330Brennus corpulentus Casey, 1897:331Brennus integer Casey, 1914:29Brennus parvulicollis Casey, 1920:176Brennus Beringi Casey, 1920:179Brennus 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 deciduousforests (e.g., Quercus), coniferous forests (e.g., Pinus, Pseudotsuga), open grassy knolls, gardens, and incultivated fields. Shaded or open areas on moderately moist ground usually covered with somevegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Adults have been found in January – July, and September –December, teneral adults in June and December. BEHAVIOR: Nocturnal, taking cover during theday under the loose bark of trees, clods of earth, fallen branches, boards, stones, and in fallentrees. 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 1924JOHNSON’S SNAIL-EATING BEETLE

Scaphinotus johnsoni Van Dyke, 1924:3Scaphinotus 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 shadedmoist 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


Scaphinotus (Scaphinotus) kelloggi (Dury) 1912KELLOGG’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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in July – August. BEHAVIOR: See description under genus, above.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollection

Scaphinotus (Nomaretus) liebecki Van Dyke 1936LIEBECK’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 havebeen 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) 1935LÖDING’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, andoccasionally 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: Occasionalclimber on cave walls while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH,UASM collections

Scaphinotus (Steniridia) lodingi obscurus (Valentine) 1935OBSCURE 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 shadedwet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking


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 NMNHcollection

Scaphinotus (Neocychrus) longiceps Van Dyke 1924LONG-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 coniferousrainforest (Sequoia, Pseudosuga) and mixed forests. MICROHABITAT: On shaded moderately moistground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. 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 inrotten tree trunks. Occasional climber on tree trunks while foraging.References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS,NMNH collectionsNotes: This species is known from midlands in northern CA, however, only one locality givenon labels is precise enough to determine altitude (that lowland one which is given above).

Scaphinotus (Scaphinotus) macrogonus Bates 1891BATES’ SNAIL-EATING BEETLE

Scaphinotus macrogonus Bates, 1891:229Scaphinotus 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 thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inSeptember. BEHAVIOR: See description under genus, above.References: Erwin (2001), Van Dyke (1938), Data from CAS, NMNH collections

Scaphinotus (Pseudonomaretus) mannii Wickham 1919MANN’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 andscrub 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: Adultshave been found in March – July, October – November, copulating pairs in June, and teneraladults in March – May. BEHAVIOR: Nocturnal, taking cover during the day in cavities under


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 NMNHcollection

Scaphinotus (Brennus) marginatus (Fischer von Waldheim) 1820MARGINED SNAIL-EATING BEETLE

Cychrus marginatus Fischer von Waldheim, 1820: pl. 7Brennus marginatus Fulleri G. Horn, 1878:179Cychrus marginatus gracilis Géhin, 1885:76Brennus cupripennis Casey, 1897:334Brennus insularis Casey, 1897:334Brennus confusus Casey, 1897:336Brennus marginatus fallax Roeschke, 1907:174Brennus columbianus Casey, 1920:180Brennus marginatus montanicus Casey, 1920:182Brennus marginatus wrangelli Casey, 1920:182Brennus 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, andcultivated fields, spruce/fir forest, Alnus thickets, in aspen scrub vegetation, snow-patches (abovetimber line), and in supratidal meadows and open coastal tundra (Aleutian Islands). Membersof 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 walkingor 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 loosebark of fallen trees, in hollow tree roots, in leaf litter and moss; gregarious (up to 50 individualsin the same shelter). Excellent climber (on fallen trees and tree-stumps) while foraging. Larvalhabitat: under stones. Adults found over wintering in fallen trees, half rotten fallen trees andtree-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 collectionsNotes: This species, as presently understood, is markedly variable across its extensive geographicand altitudinal range. It is ripe for an intense genomics study.

Scaphinotus (Pseudonomaretus) merkelii (G. Horn) 1890MERKEL’S SNAIL-EATING BEETLE

Sphaeroderus Merkelii G. Horn, 1890:71Cychrus idahoensis Webb, 1901:133


Distribution: Native, New World. CANADA – BC; USA – ID, MT.Way of Life: MACROHABITAT: Lowlands to uplands, 700 – 1675 meters altitude, in mixedforests. MICROHABITAT: On shaded rather dry ground. DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave 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 1882MEXICAN 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 towalking 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) 1868OBLIQUE SNAIL-EATING BEETLE

Cychrus obliquus LeConte, 1868:61Brennus convergens Casey 1897:326Brennus opacicollis Casey 1897:327Brennus 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 thusvagility 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 NMNHcollection

Scaphinotus (Brennus) oreophilus (Rivers) 1890MOUNTAIN LOVING SNAIL-EATING BEETLE

Cychrus oreophilus Rivers, 1890:111Scaphinotus oreophilus Hoppingi Roeschke, 1907:183Brennus 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


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 1988OZARK 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. Seedescription under genus, above.References: Erwin (2001), Larochelle & Larivière (2003), Will (pers. comm.)

Scaphinotus (Scaphinotus) petersi biedermani Roeschke 1907BIEDERMAN’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 coniferousforests. MICROHABITAT: On shaded moist ground on slopes. DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave been found in June – September. BEHAVIOR: Nocturnal, taking cover in the day understones and bits of fallen trees.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollection

Scaphinotus (Scaphinotus) petersi catalinae Van Dyke 1924CATALINA 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 flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inJune – 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


Scaphinotus (Scaphinotus) petersi corvus (Fall) 1910RAVEN 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, coniferousforests. 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 coverin 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 1938GRAHAM’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 coniferousand mixed 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 – September. BEHAVIOR: Nocturnal, taking coverin the day under stones and bits of fallen trees.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollection

Scaphinotus (Scaphinotus) petersi kathleenae Ball 1966KAY’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 ponderosaDouglas), and Gambel’s Oak (Quercus gambelii Nuttall). MICROHABITAT: On shaded moist, butwell-drained, loamy ground underlain by coarse gravel, particularly on steep slopes. DISPERSAL


OCCURRENCE: Adults have been found in August. BEHAVIOR: Nocturnal, taking cover in the dayunder fallen trees, bits of fallen trees, and in leaf litter.References: Ball, (1966), Erwin (2001), Larochelle & Larivière (2003), Data from NMNH,UASM collections


Scaphinotus (Scaphinotus) petersi petersi Roeschke 1907PETERS’ 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, coniferousforest. MICROHABITAT: On shaded moist ground. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in June – August. BEHAVIOR: Nocturnal, taking cover in the day under fallen trees andbits of fallen trees.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollection

Scaphinotus (Brennus) punctatus (LeConte) 1859PUNCTATE SNAIL-EATING BEETLE

Cychrus punctatus LeConte, 1859:69Cychrus mimus G. Horn, 1874:20Brennus gravidus Casey, 1897:317Brennus 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 vagilitylimited 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 theday under stones, fallen trees, and dead leaves. Gregarious. Adults eat snails, live and deadcaterpillars, berries, and other beetles.References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Noonan (1967),Data from NMNH collection

Scaphinotus (Pseudonomaretus) regularis (LeConte) 1884MEASURED 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 thusvagility 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 NMNHcollection


Scaphinotus (Pseudonomaretus) relictus (G. Horn) 1881RILICTUAL 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 coverduring 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 1935RIDINGS’ 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 vagilitylimited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Nocturnal, takingcover during the day.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935)

Scaphinotus (Steniridia) ridingsii monongahelae Leng 1917MONONGAHELA SNAIL–EATING BEETLE

Scaphinotus monongahelae Leng, 1917:36Irichroa 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. Moderaterunner. SEASONAL OCCURRENCE: Adults have been found in May – October, teneral adults inmid-July. BEHAVIOR: Nocturnal, taking cover during the day under fallen trees. Occasionalclimber (on tree trunks) while foraging.References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection

Scaphinotus (Steniridia) ridingsii ridingsii (Bland) 1863RIDINGS’ SNAIL-EATING BEETLE

Cychrus Ridingsii Bland, 1863:353

Distribution: Native, New World. USA – VA.


Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 915 meters altitude, forests inriparian habitats. MICROHABITAT: Found in rocky, moist sites in the shade along streams andin 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 arepredaceous on snails; they have been observed mating in September. Frequent climber onstones and boulders while foraging.References: Erwin (1981, 2001), Larochelle & Larivière (2003), Valentine (1935), Data fromNMNH collection

Scaphinotus (Brennus) riversi (Roeschke) 1907RIVERS’ 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults havebeen found in May – October. BEHAVIOR: Nocturnal, taking cover during the day understones and fallen trees.References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS,NMNH collections

Scaphinotus (Brennus) rugiceps incipiens (Casey) 1897CASEY’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 andconiferous forest MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound 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) 1872WRINKLE-HEADED SNAIL-EATING BEETLE

Cychrus rugiceps G. Horn, 1872:143Brennus porcatus Casey, 1897:328Brennus compositus Casey, 1897:332Brennus rugiceps congener Casey, 1914:28

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


Way of Life: MACROHABITAT: Lowlands to uplands, 75 – 1880 meters altitude, mixed andconiferous 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) 1913SCHWARZ’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 grassybalds. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. Moderate runner. Strong burrower. SEASONAL OCCURRENCE: Adults have been foundin June, and August – September. BEHAVIOR: Nocturnal, taking cover during the day in leaflitter, 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 1938ROESCHKE’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 coniferousforests. MICROHABITAT: Shaded areas on moist ground, along brooks and near springs. DISPERSAL


OCCURRENCE: Adults have been found in July. BEHAVIOR: Nocturnal, taking cover during theday under stones and fallen trees.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection

Scaphinotus (Scaphinotus) snowi roeschkei Van Dyke 1907ROESCHKE’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: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollection


Scaphinotus (Scaphinotus) snowi snowi (LeConte) 1881SNOW’S SNAIL-EATING BEETLE

Cychrus Snowi LeConte, 1881:71Scaphinotus 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 coniferousforests. MICROHABITAT: Shaded areas on moist ground, along brooks and near springs. DISPERSAL


OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Nocturnal, taking coverduring the day under stones and fallen trees.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollection

Scaphinotus (Brennus) striatopunctatus (Chaudoir) 1844PORE-WINGED SNAIL-EATING BEETLE

Cychrus striatopunctatus Chaudoir, 1844:476Brennus decipiens Casey, 1897:316Brennus subdepressus Casey, 1920:177

Distribution: Native, New World. USA – CA.Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 1524 meters altitude, mostly indeciduous forests (e.g., Quercus), but also in canyons and on grassy knolls. MICROHABITAT: Shadedor open areas on moderately moist, sandy ground. DISPERSAL ABILITIES: Brachypterous, henceflightless 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 offallen 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) 1863SLENDER 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 ofwater. MICROHABITAT: Adults are found on wet ground. DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave been found in January – August, and October – December. BEHAVIOR: Nocturnal, takingcover during the day.References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS,NMNH, UASM collections


Scaphinotus (Maronetus) tenuis (Casey) 1914THIN 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) 1914THREE-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


OCCURRENCE: Adults have been found in April – October. BEHAVIOR: Nocturnal, taking coverduring 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) 1787ONE-COLOR SNAIL-EATING BEETLE

Cychrus unicolor Fabricius, 1787:198Scaphinotus heros T.W. Harris, 1839:196, new synonymyScaphinotus grandis Gistel, 1857:52, n. dubiumScaphinotus Hunteri Crotch, 1873:5Scaphinotus Shoemakeri Leng, 1914:143, new synonymyScaphinotus 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, oftenalong streams and near springs. MICROHABITAT: Found in and around rocky outcroppings inthe eastern deciduous forest, usually under fallen trees or at night foraging in the leaf litter andshaded areas with wet, rocky ground. DISPERSAL ABILITIES: Brachypterous, hence flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary – November. BEHAVIOR: Nocturnal, taking cover during the day under stones, fallentrees, and dead leaves. Adults and larvae eat snails, at least in part, but probably also take other


kinds of food, such as white grubs of the May (June) Beetle (Phyllophaga spp.). Adults and larvaeprobably 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 NMNHcollection

Scaphinotus (Maronetus) unistriatus (Darlington) 1931ONE-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 alongslopes. 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 1907VAN DYKE’S SNAIL-EATING BEETLE

Scaphinotus Van Dykei Roeschke, 1907:136Scaphinotus fuchsi Roeschke, 1907:570

Distribution: Native, New World. USA – AZ.Way of Life: MACROHABITAT: Uplands to mountains, 1830 – 2440 meters altitude, in coniferousforests, 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 coverduring the day under stones and bits of fallen trees.References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNHcollection

Scaphinotus (Stenocantharis) velutinus (Ménétriés) 1843VELVET SNAIL-EATING BEETLE

Cychrus velutinus Ménétriés, 1843:53Pemphus longipes Casey, 1897:339Pemphus opacus Casey, 1899:97

Distribution: Native, New World. USA – CA, OR.Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1036 meters altitude, in matureconiferous 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


Scaphinotus (Brennus) ventricosus (Dejean) 1831POTBELLIED SNAIL-EATING BEETLE

Cychrus Ventricosus Dejean, 1831:527Cychrus lativentris Motschulsky, 1850:358Cychrus ovalis Motschulsky, 1859:162Cychrus alternatus Motschulsky, 1859:163Cychrus fuchsianus Rivers, 1890:71Brennus symmetricus Casey, 1897:319Brennus strictus Casey, 1897:322Brennus brevicollis Casey, 1920:178

Distribution: Native, New World. USA – CA, OR.Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 515 meters altitude, in mostlydeciduous forests (e.g., Quercus) and poison-oak thickets, as well as on open grassy knolls andedges of reservoirs. MICROHABITAT: Shaded and open ground areas on moderately moist ground,sometimes sandy soils. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limitedto walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have been found inJanuary – 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 NMNHcollection

Scaphinotus (Scaphinotus) viduus (Dejean) 1826BEREFT SNAIL-EATING BEETLE

Cychrus Viduus Dejean, 1826:12Cychrus Leonardii T.W. Harris, 1839:193Cychrus 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 coniferousforests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). At lower altitudes in mixed and deciduousforests (e.g., Quercus spp., Acer spicatum Lam.) and adjacent fields. MICROHABITAT: On shadedwet 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, teneraladults 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 underthe loose bark of trees, and in small cells dug under fallen trees, fallen tree-stumps, and piecesof wood. Adult are predaceous on snails, slugs, and caterpillars. Larvae over winter under fallentrees; they are gregarious.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection


Scaphinotus (Steniridia) violaceus carolinae (Valentine) 1935CAROLINA 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 thusvagility 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) 1863VIOLET 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 havebeen found in May – July, and September – November. BEHAVIOR: Nocturnal, taking coverduring 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 1959WEBB’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


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 1826FALSE 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


Type species: Cychrus stenostomus Weber, 1801Distribution: Nearctic.Habitat: According to Lindroth (1961), species in this genus hibernate as adults, unlike mostother Cychrini. They are inhabitants of mesic forests.References: Erwin (2001), Lindroth (1961)

Sphaeroderus bicarinatus (LeConte) 1853TWO-RIDGED FALSE SNAIL-EATING BEETLE

Cychrus bicarinatus LeConte, 1853:399Sphaeroderus 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 coniferousforests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir, (at lower altitudes in mixed and deciduousforests, e.g., Quercus spp., Acer spicatum Lam.). MICROHABITAT: Shaded areas on moderately moistground covered with deep leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thusvagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inMarch – October, and December, teneral adults in late August. BEHAVIOR: Nocturnal, takingcover during the day in leaf litter, under the loose bark of fallen trees, in and under rotten fallentrees, under stones, and under moss carpets.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

Sphaeroderus canadensis canadensis Chaudoir 1861CANADIAN FALSE SNAIL-EATING BEETLE

Sphaeroderus canadensis Chaudoir, 1861:498Sphaeroderus palpalis Motschulsky, 1866:312Sphaeroderus 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: Shadedareas 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 mosscarpets. Larval habitat similar to adult, in leaf litter. Coitus lasting 101 – 142 minutes. Adultshave been found over wintering on higher and drier ground, at the summit of wooded slopesand 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 collectionNotes: Gidaspow (1973) regarded this name as in the genus Nomaretus.


Sphaeroderus canadensis lengi Darlington 1933LENG’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 coniferousforests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir, (at lower altitudes in mixed and deciduousforests, e.g., Quercus spp., Acer spicatum Lam.). MICROHABITAT: Shaded areas on moderately moistground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. 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) 1910INDIANA 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 flightlessthus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found inFebruary, 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 1848BREVOORT’S FALSE SNAIL-EATING BEETLE

Sphaeroderus Brevoorti LeConte, 1848:343Sphaeroderus 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 metersaltitude, in coniferous and mixed forests or at their edges. Adults occur along rivers, brooks, atthe seashore, and on slopes. MICROHABITAT: Shaded area on wet or moist ground covered withdeep leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking 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 theday 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. Accordingto Larochelle & Larivière (2003), “adults have been found over wintering on higher and drierground, in woods; under moss and in leaf litter, at the base of tree-stumps and trees, under theloose bark of fallen trees, in fallen rotten branches, and under stones.” Adult and larvae arepredaceous on snails. Coitus has been recorded as lasting up to 72 hours.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection


Sphaeroderus nitidicollis nitidicollis Guérin-Méneville 1829POLISHED-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, indeciduous, mixed, or coniferous forests, and in thickets. Adults occur along rivers, brooks, atthe seashore. MICROHABITAT: Shaded areas on wet or moist ground covered with deep leaflitter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking orrunning. SEASONAL OCCURRENCE: Adults have been found in June – September; copulatingpairs seen in July; teneral adults in July – August. BEHAVIOR: Nocturnal, taking cover during theday 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 22hours. Adults hibernate.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

Sphaeroderus nitidicollis schaumii Chaudoir 1861SCHAUM’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 deciduousforests (e.g., Acer; Celtis). MICROHABITAT: Shaded areas on moderately moist ground. DISPERSAL


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 understones. Adult are predaceous on snails.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

Sphaeroderus stenostomus lecontei Dejean 1826LECONTE’S FALSE SNAIL-EATING BEETLE

Sphaeroderus Lecontei Dejean, 1826:15Sphaeroderus niagarensis Laporte de Castelnau, 1833:390Sphaeroderus 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 – 1067meters 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 leaflitter, 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.


SEASONAL OCCURRENCE: Adults have been found in January – December; copulating pairs inApril – July, September – November; ovipositing females in May – July; late instar larvae inAugust; teneral adults in July – September. BEHAVIOR: Mostly nocturnal, taking cover duringthe day in leaf litter, under fallen trees and stones, in mosses, or under the loose bark of fallentrees; 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 onhigher and drier ground, in woods and at their edges, in leaf litter, in rotten tree-stumps andfallen trees, under clods of earth, and under stones. Adult are predaceous on snails, slugs, andlepidopterous caterpillars. Occasional climber while foraging. Coitus has been recorded as lastingup to 72 hours.References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1955), Data from NMNHcollection

Sphaeroderus stenostomus stenostomus (Weber) 1801NARROW-MOUTHED FALSE SNAIL-EATING BEETLE

Cychrus stenostomus Weber, 1801:43Sphaeroderus 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 deciduousand coniferous forests and at the borders of eutrophic marshes. MICROHABITAT: Shaded areason dry ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited towalking or running. SEASONAL OCCURRENCE: Adults have been found in February – November;copulating pairs in April, October – November; teneral adults in July. BEHAVIOR: MostlyNocturnal, taking cover during the day under dead leaves, stones, and fallen trees, sometimesactive on cloudy days. Adults hibernate. Adult are predaceous on snails and slugs.References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collectionNotes: Casey’s single type specimen from PA (NMNH) matches perfectly one of three specimenscollected together on the Blue Ridge summit in Franklin County. The only differences betweenCasey’ specimen and the matching one and the other two specimens are that the pronotal baseis not extensively punctulate and the elytra are a bit more parallel-sided. The other two specimensare perfectly good S. s. stenomus (Weber). In a series of 298 specimens in NMNH from throughoutthe range of S. s. stenomus (Weber), both these attributes are variable. Since the variation existsin co-collected specimens, subspecies status is not possible.

CICINDELITAECollyridini 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 1835Geographic status: Neotropical.References: Erwin (2001), Lorenz (2005), Naviaux (1998)


Ctenostoma Klug 1821COMB-MOUTHED BEETLES

(Figure 22)

Ctenostoma Klug, 1821:302Caris, Fisher von Waldheim, 1821:99Procephalus 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 thata new name is needed for this subgenus of species.

Naviauxiana Erwin, new replacement name for Procephalus Laporte de Castelnau, 1834 [valid subgenusfor 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 1835Type 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 ofthe Mata Atlântica; some altitude records associated with the Andes and Central America indicatethey also occur in cloud forests. These long-legged beetles, many of which are ant-shaped andbrachypterous, are found on twigs and foliage in the canopy and understory of moist tropicalforests, rain forests, and likely cloud forests; also adults have been found in fogging samples ofthe tall river-side grass, Gynerium sagittatum (Aubl.) Beauv and stands of the bamboo, Guaduaweberbaueri Pilger. Larvae use holes in rotting twigs and perhaps bamboo culms much as othertiger beetle larvae use holes in the ground. Some species make their burrows in the end ofbroken twigs, others in the lateral surfaces of the twig. The burrows are enlarged as the larvagrows. Both adults and larvae are predatory on insects. Although Zikan (1929) reportedobservations on the way of life of some species in South America, particularly Brazil, the genusremains inadequately known. Naviaux (1998) noted that adults of some species seem to mimiccerambycid beetles; perhaps it is the other way round since all carabids possess well-developedchemical 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 1998SHORT 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


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

Ctenostoma (Ctenostoma) acciavatti Naviaux 1998ACCIAVATTI’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, probablycapable 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 1998BRASSY 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 associationwith the plant, Ocotea whitei Woodson. DISPERSAL ABILITIES: Macropterous, probably capable offlight. SEASONAL OCCURRENCE: Adults have been found in March – June. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998), Data from NMNH collection

Ctenostoma (Myrmecilla) agnatum agnatum Chaudoir 1860SMALL-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 rainforestand cloud forests. MICROHABITAT: Canopy, closed, at an average of 37 meters height. Also, insuspended 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 vagilitylimited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February –March, June, and September – October. BEHAVIOR: See description under genus, above. Manyexamples 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. Adultsof 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


Ctenostoma (Myrmecilla) agnatum fuscum Naviaux 1998SOMBER 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


OCCURRENCE: The adult holotype was found in October. BEHAVIOR: See description undergenus, above.References: Erwin (2001), Naviaux (1998), Data from NMNH collection

Ctenostoma (Myrmecilla) albofasciatum Chaudoir 1850WHITE-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, henceflightless 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 1925NARROWLY-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 inMarch – June, and August. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Data from NMNH collection

Ctenostoma (Naviauxiana) arnaudi Naviaux 1998ARNAUD’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 andcloud 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)


Note: Pearson et al. (1999a) cast doubt on the western slope 650 meter altitude record in EsmeraldasProvince believing this species is an Amazon Basin one, thus not occurring in the west.

Ctenostoma (Myrmecilla) asperulum Bates 1868ROUGH COMB-MOUTHED BEETLE

Ctenostoma asperulum Bates, 1868:278


Ctenostoma (Ctenostoma) bahiaense Naviaux 1998BAHIA 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 descriptionunder genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Ctenostoma) batesii Chaudoir 1860BATES’ 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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in February, June, andOctober. BEHAVIOR: See description under genus, above. Many examples were obtained fromcanopy 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 1860DOUBLE-CRESTED COMB-MOUTHED BEETLE

Ctenostoma bicristatum Chaudoir, 1860:282

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, henceflightless 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 1831DOUBLE-MARKED COMB-MOUTHED BEETLE

Ctenostoma Bifasciatum Dejean, 1831:272


Ctenostoma (Euctenostoma) bifasciatum horni Naviaux 1998HORN’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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Euctenostoma) bondari W. Horn 1939BONDAR’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, henceflightless 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 1998BRENDELL’S COMB-MOUTHED BEETLE

Ctenostoma brendelli Naviaux, 1998:80

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


Way of Life: MACROHABITAT: Lowlands, 133 meters altitude, in dry forest. MICROHABITAT:Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005)

Ctenostoma (Myrmecilla) brevicorne W. Horn 1898SHORT-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 foliagein the understory of moist tropical forest types, and larvae are found in burrows in twigs andsmall 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 1931SHORT-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, probablycapable 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 1837MANNERHEIM’S COMB-MOUTHED BEETLE

Ctenostoma breviusculum von Mannerheim, 1837:20



Ctenostoma (Neoprocephalus) brunneum Naviaux 1998BROWNISH 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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in April. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005)

Ctenostoma (Neoprocephalus) cassolai Naviaux 1998CASSOLA’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, probablycapable 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 1998CAYENNE 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, probablycapable 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) 1895CHAUDOIR’S COMB-MOUTHED BEETLE

Procephalus Chaudoiri W. Horn, 1895:353Ctenostoma Chaudoiri Gounelli W. Horn, 1898:27Ctenostoma 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 ofthe Tropical Moist Forest of Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT:Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably


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 1998COMPACT 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 towalking or running. SEASONAL OCCURRENCE: Adults have been found in May – August, andOctober. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Data from NMNH collection

Ctenostoma (Myrmecilla) coracinum Naviaux 1998BLACKISH 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 1868LITTLE-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 offlight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Myrmecilla) crucifrons W. Horn 1911CROSS-FACED COMB-MOUTHED BEETLE

Ctenostoma crucifrons W. Horn, 1911:154

Distribution: Native, New World. PERÚ.


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 descriptionunder genus, above.References: Erwin (2001), Naviaux (1998), Data from CAS collection

Ctenostoma (Neoprocephalus) cylindratum Naviaux 1998ROLLER-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, probablycapable 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 1998DAVIDSON’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 rainforestand tropical dry forest. MICROHABITAT: Arboreal, on twigs and leaves above the ground.DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adultshave been found in June – July. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005)

Ctenostoma (Myrmecilla) dentifrons W. Horn 1901TOOTH-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 wascollected in the state of Espirito Santo, Brazil, the Mata Atlântica in Brazil was still growing. Atpresent, 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)


Ctenostoma (Neoprocephalus) deuvei Naviaux 1998DEUVE’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 beenfound 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 1898DOKHTUROW’S COMB-MOUTHED BEETLE

Ctenostoma Dokhturowi W. Horn, 1898:32


Ctenostoma (Naviauxiana) dormeri W. Horn 1898DORMER’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 rainforestand cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL

ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have beenfound 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 1998DURANTON’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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in February – July, andDecember. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Naviauxiana) ebenina Bates 1868EBONY 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, andOctober – November. BEHAVIOR: See description under genus, above. Adults are attracted tolights at night.References: Erwin (2001), Naviaux (1998), Pearson, et al. (1999b), Data from D. Pearsoncollection

Ctenostoma (Euctenostoma) eburatum Bates 1872IVORY 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: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Naviauxiana) ecuadorense Naviaux 1998ECUADORIAN 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 rainforestand cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL

ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found inJanuary – 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 1998ERWIN’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 rainforestand cloud forests. MICROHABITAT: Adults are found on twigs and foliage in the shadyunderstory of moist tropical forest types, and larvae are found in burrows in twigs and small


branches. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE:Adults have been found in April – July. BEHAVIOR: Adults are predaceous on small arborealarthropods, seeking them on twigs and leaves of forest trees and shrubs. Adults occasionallyfound at lights at night.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Data from NMNHcollection

Ctenostoma (Euctenostoma) ferum Naviaux 2005WILD COMB-MOUTHED BEETLE

Ctenostoma ferum Naviaux, 2005:267

Distribution: Native, New World. BRAZIL.Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves abovethe ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walkingor running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above.References: Naviaux (2005)

Ctenostoma (Myrmecilla) flexuosum Naviaux 1998FLEXED-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, henceflightless 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) 1801ANT-LIKE COMB-MOUTHED BEETLE

Collyris formicarium Fabricius, 1801:226Ctenostoma Jacquieri Dejean, 1831:271Ctenostoma 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 inNovember. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Data from CAS collection


Ctenostoma (Euctenostoma) fryi Chaudoir 1865FRY’S COMB-MOUTHED BEETLE

Ctenostoma Fryi Chaudoir, 1865:49


Ctenostoma (Myrmecilla) gautardi Chaudoir 1869GAUTARD’S COMB-MOUTHED BEETLE

Ctenostoma Gautardi Chaudoir, 1869:22


Ctenostoma (Neoprocephalus) germaini W. Horn 1902GERMAIN’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: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998), Pearson, et al. (1999b)

Ctenostoma (Myrmecilla) globifrons W. Horn 1898SPHERICAL-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 ofthe tropical moist forest in Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT:Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in May – June. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Neoprocephalus) guatemalense Brouerius van Nidek 1960GUATEMALAN 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: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Ctenostoma) guyanense W. Horn 1998GUIANAN 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, probablycapable 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 1894HEYDEN’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 abovethe ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL


Ctenostoma (Myrmecilla) hirsutum W. Horn 1892HAIRY COMB-MOUTHED BEETLE

Ctenostoma hirsutum W. Horn, 1892:46Ctenostoma macilentum Klug, 1898:29.



Ctenostoma (Neoprocephalus) hovorei Naviaux & Brzoska 2005PACO’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, probablycapable 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 1880DOHRN’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 abovethe ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL


Ctenostoma (Neoprocephalus) ibidion jolyi Naviaux 1998JOLY’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 cloudforests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inMay – July, and October. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005)

Ctenostoma (Myrmecilla) ichneumoneum ellipticum Naviaux 1998ELLIPSUS-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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January – March, and November – December. BEHAVIOR: See description undergenus, above.References: Erwin (2001), Naviaux (1998, 2002)


Ctenostoma (Myrmecilla) ichneumoneum ichneumoneum Dejean 1826WASP-LIKE COMB-MOUTHED BEETLE

Ctenostoma Ichneumoneum Dejean, 1826:436Ctenostoma 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January – February and November – December. BEHAVIOR: See description undergenus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Neoprocephalus) immaculatum W. Horn 1925NON-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 offlight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Euctenostoma) inca Naviaux 1998INCA 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 rainforestand cloud forests. MICROHABITAT: Dry hanging fronds of Astrocaryum palm in the upperfloodplain forest on the Rio Manu. DISPERSAL ABILITIES: Macropterous, probably capable offlight. 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 NMNHcollection

Ctenostoma (Myrmecilla) infimum Naviaux 1998LOWLY 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


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

Ctenostoma (Naviauxiana) insigne Chaudoir 1860MARKED 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, probablycapable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998, 2002)

Ctenostoma (Neoprocephalus) intermedium Naviaux 1998INTERMEDIATE COMB-MOUTHED BEETLE

Ctenostoma intermedium Naviaux, 1998:71

Distribution: Native, New World. VENEZUELA.Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves abovethe ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL


Ctenostoma (Ctenostoma) jekelii jekelii Chevrolat 1858JEKEL’S COMB-MOUTHED BEETLE

Ctenostoma formicarum, [sic] Fabricius: Klug, 1821:304Ctenostoma Jekelii Chevrolat, 1858:484Ctenostoma Jekeli W. Horn, 1898:28

Distribution: Native, New World. BRAZIL; FRENCH GUIANA.Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in rainforest. MICROHABITAT: Adultsare found on twigs and foliage in the shady understory of moist tropical forest types, and larvaeare found in burrows in twigs and small branches. DISPERSAL ABILITIES: Brachypterous, winglength 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 smallarboreal arthropods, seeking them on twigs and leaves of forest trees and shrubs.References: Erwin (2001), Naviaux (1998), Data from D. Pearson collection


Ctenostoma (Ctenostoma) jekelii raresculptum W. Horn 1898RARE-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 partof the tropical moist forest in Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT:Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, winglength variable, however adults are flightless, thus vagility limited to walking or running. SEASONAL

OCCURRENCE: Adults have been found in January – March. BEHAVIOR: See description undergenus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Neoprocephalus) johnsoni Naviaux 1998JOHNSON’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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Euctenostoma) klugeanum W. Horn 1915KLUG’S COMB-MOUTHED BEETLE

Ctenostoma Klugeanum W. Horn, 1915:424Ctenostoma Klugi W. Horn, 1898:28


Ctenostoma (Neoprocephalus) laeticolor Bates 1878JOYFUL-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


capable of flight. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Neoprocephalus) landolti Steinheil 1877LANDOLT’S COMB-MOUTHED BEETLE

Ctenostoma Landolti Steinheil, 1877:48


Ctenostoma (Naviauxiana) longipalpe Naviaux 1998LONG-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 cloudforests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inMay – 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 1860MOURNFUL 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, capableof flight. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: See descriptionunder genus, above. One example was obtained from canopy foggings in southeastern Perú, inwet to dry transitional season. Adults are attracted to lights.References: Erwin (2001), Erwin (field notes), Naviaux (1998), Pearson, et al. (1999b), Datafrom D. Pearson, NMNH collections


Ctenostoma (Myrmecilla) luteum Naviaux 1998YELLOWISH 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 limitedto 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 1834THIN COMB-MOUTHED BEETLE

Ctenostoma macilentum Klug, 1834:42


Ctenostoma (Neoprocephalus) maculicorne (Chevrolat) 1856SPOTTED-HORN COMB-MOUTHED BEETLE

Procephalus maculicornis Chevrolat, 1856:352Ctenostoma 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 rainforestand cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL

ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found inApril – July, and October – November. BEHAVIOR: See description under genus, above. Adultshave 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 1998SMALL-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


of flight. SEASONAL OCCURRENCE: Adults have been found in January – March. BEHAVIOR: Seedescription under genus, above. Adults occasionally found at lights at night.References: Erwin (2001), Naviaux (1998), Data from NMNH collection

Ctenostoma (Euctenostoma) magnum Naviaux 1998VERY 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, probablycapable 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) 1834METALLIC 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 abovethe ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL


Ctenostoma (Myrmecilla) minusculum Naviaux 2002VERY 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in November. BEHAVIOR: See description under genus, above.References: Naviaux (1998)

Ctenostoma (Myrmecilla) modicum Naviaux 1998MODEST 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:


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 2002MONNE’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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown.BEHAVIOR: See description under genus, above.References: Naviaux (2002)

Ctenostoma (Naviauxiana) nigrum lecourti Naviaux 1998LECOURT’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 rainforestand cloud forests, Yungas. MICROHABITAT: Suspended dry leaves. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inMarch – April, and October – November. BEHAVIOR: See description under genus, above. Anadult was found in suspended dry leaves in a stand of bamboo (Guadua weberbaueri Pilger) about2.5 meters off the ground using insecticidal fogging techniques.References: Erwin (2001), Erwin (field notes), Naviaux (1998), Pearson, et al. (1999b), Datafrom NMNH collection

Ctenostoma (Naviauxiana) nigrum nigrum Chaudoir 1860BLACK 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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Neoprocephalus) nitidum Naviaux 1998BRILLIANT 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 rainforestand cloudforests, Yungas. MICROHABITAT: Arboreal, on twigs and leaves above the ground.DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adultshave 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 standof 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 1860SLOPED 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 4mabove the ground and in the mid-strata of the rainforest. DISPERSAL ABILITIES: Brachypterous,hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adultshave been found in October – November. BEHAVIOR: See description under genus, above. Anadult from Perú was collected in a malaise trap, however the specimen is brachypterous. Adultswere found in the microhabitat described above using insecticidal fogging techniques.References: Erwin (2001), Erwin (field notes), Naviaux (1998), Pearson, et al. (1999b), Datafrom D. Pearson, NMNH collection

Ctenostoma (Myrmecilla) oblitum Chaudoir 1860EFFACED 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in December. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Naviauxiana) onorei colombiense Naviaux 1998COLOMBIAN 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 inJanuary – March. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Naviauxiana) onorei onorei Naviaux 1998ONORE’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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in April. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Naviauxiana) ornatum ornatum Klug 1834ORNATE 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 offlight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Naviauxiana) ornatum werneri Navaiux 1998WERNER’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 metersaltitude. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inJanuary – February, and November. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Naviauxiana) panamense Naviaux & Brzoska 2005PANAMANIAN 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, probablycapable 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 1998PARALLEL-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 descriptionunder genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Myrmecilla) parvulum Navaiux 1998DIM 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in November and December. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Naviauxiana) pearsoni Navaiux 1998PEARSON’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, capableof flight. SEASONAL OCCURRENCE: Adults have been found in April – July. BEHAVIOR: Seedescription under genus, above. Adults are attracted to lights at night.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Myrmecilla) plicaticolle W. Horn 1911PLEATED-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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in October – December. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Microprocephalus) pusillum minimum Navaiux 1998TINY-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: Adultshave been found in October. BEHAVIOR: See description under genus, above. Adults have beencommonly found using insecticidal fogging techniques.References: Erwin (2001), Erwin et al. (2004), Naviaux (1998), Pearson et al. (1999a), Datafrom NMNH collection

Ctenostoma (Microprocephalus) pusillum pusillum Navaiux 1998TINY 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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in May, July – August, andDecember. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Myrmecilla) pygmaeum concolor Naviaux 1998UNIFORM 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January and December. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Myrmecilla) pygmaeum pygmaeum (Lacordaire) 1843PIGMY COMB-MOUTHED BEETLE

Myrmecilla pygmaea Lacordaire, 1843:120


Ctenostoma (Euctenostoma) rapillyi Naviaux 1998RAPILLY’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, probablycapable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus,above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Euctenostoma) regium Naviaux 1998REGAL 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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in November – December.BEHAVIOR: See description under genus, above. According to Pearson, et al. (1999a), “Unlikealmost all other species of this genus in Ecuador, the eyes are not distinctively wider than therest of the head. The pronotum is distinctly rounded and domed. The apex of each elytron hasa shallow scallop-shaped notch. The majority of these peculiar characters are apparently toenhance mimicry of the powerfully stinging ant Paraponera [clavata (Fabricius) 1775)]. Not onlyis this beetle the same size and shape as this large ant, but it walks and behaves similarly, even tothe 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 ofprimary forest in the lowlands of eastern Ecuador and Peru.”References: Erwin (2001), Naviaux (1998), Pearson et al. (1999a), Data from D. Pearsoncollection


Ctenostoma (Myrmecilla) rivalieri Naviaux 1998RIVALIER’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 1904ROUGH-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 beenfound 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 1913FALSE 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, probablycapable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus,above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Ctenostoma) rugiferum rugiferum (W. Horn) 1895WRINKLED COMB-MOUTHED BEETLE

Procephalus rugifer W. Horn, 1895:354



Ctenostoma (Euctenostoma) rugosum Klug 1824FOLDED COMB-MOUTHED BEETLE

Ctenostoma rugosum Klug, 1824:7Ctenostoma 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, winglength 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 1860SAHLBERG’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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January and December. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Naviauxiana) sallei Chaudoir 1860SALLE’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 abovethe ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL


Ctenostoma (Myrmecilla) schaumi pilosulum W. Horn 1898FELT-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 ofthe tropical moist forest in Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT:Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in January – March. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)


Ctenostoma (Myrmecilla) schaumi schaumi W. Horn 1895SCHAUM’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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in November. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005)

Ctenostoma (Myrmecilla) schmalzi paraguayense Brouerius van Nidek 1956PARAGUAYAN 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in February. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Myrmecilla) schmalzi schmalzi W. Horn 1898SCHMALZ’S COMB-MOUTHED BEETLE

Ctenostoma Schmalzi W. Horn, 1898:32


Ctenostoma (Naviauxiana) simile Naviaux 1998VERY 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 cloudforests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES:Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found inMarch and June. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Data from NMNH collection


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

Ctenostoma simpliceps W. Horn, 1900:361


Ctenostoma (Naviauxiana) spinosum Naviaux 1998SPINY 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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in May. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Naviauxiana) subtilesculptum W. Horn 1913SUBTLY-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 dryforest, 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) 1834VERY 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 rainforestand cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL

ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have beenfound in March and November. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Pearson et al. (1999a)


Ctenostoma (Euctenostoma) sumlini Naviaux 1998SUMLIN’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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in September. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Ctenostoma) transversum Naviaux 1998TRANSVERSELY-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, probablycapable 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) 1821THREE-SPOTTED COMB-MOUTHED BEETLE

Caris fasciatum Fischer von Waldheim, 1821: pl. 1Caris trinotata Fischer von Waldheim, 1821:99Ctenostoma Fischeri Chaudoir, 1848:22


Ctenostoma (Neoprocephalus) tumidum Naviaux 1998SWOLLEN-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, probablycapable of flight. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Seedescription under genus, above.References: Erwin (2001), Naviaux (1998), Data from NMNH collection


Ctenostoma (Neoprocephalus) turnbowi Naviaux 1998TURNBOW’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 inMay. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Euctenostoma) tyrannum (Thomson) 1859MASTER COMB-MOUTHED BEETLE

Procephalus tyrannus Thomson, 1859:92


Ctenostoma (Myrmecilla) unifasciatum affine W. Horn 1894KIN COMB-MOUTHED BEETLE

Ctenostoma affine W. Horn, 1894:109Ctenostoma Gautardi affine W. Horn, 1908:91


Ctenostoma (Myrmecilla) unifasciatum unifasciatum Dejean 1831ONE-LINED COMB-MOUTHED BEETLE

Ctenostoma trinotatum Klug, 1824:5Ctenostoma Unifasciatum Dejean, 1831:272Ctenostoma trifasciatum Lacordaire, 1843:118Ctenostoma Klugii Lacordaire, 1843:119Ctenostoma laceratum R.F. Sahlberg, 1844:503

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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have beenfound in April and November. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Neoprocephalus) vairai Cassola 2001VAIRA’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 inFebruary. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Myrmecilla) vicinum Naviaux 1998NEIGHBORHOOD COMB-MOUTHED BEETLE

Ctenostoma vicinum Naviaux, 1998:136


Ctenostoma (Neoprocephalus) wappesi Naviaux 1998WAPPES’ 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 rainforestand cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL

ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have beenfound in May – July. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Data from CAS collection


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

Ctenostoma wiesneri Naviaux, 1998:114


Ctenostoma (Myrmecilla) zerchei Naviaux 1998ZERCHE’S COMB-MOUTHED BEETLE

Ctenostoma zerchei Naviaux, 1998:131


Ctenostoma (Salvioides) zikani W. Horn 1911ZIKAN’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 inJanuary and December. BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

Ctenostoma (Myrmecilla) zonata Chaudoir 1860GIRDLED 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, henceflightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown.BEHAVIOR: See description under genus, above.References: Erwin (2001), Naviaux (1998)

GENERAL BIBLIOGRAPHY FOR CARABOIDEA OF THE WESTERN HEMISPHERE | 207

General Bibliography for Caraboidea of theWestern Hemisphere

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BALL, G.E. (1978) The species of the Neotropical Genus Trichopselaphus Chaudoir (Coleoptera:Carabidae: Harpalini): Classification, phylogeny and zoogeography. Quaestiones entomolgicae,14, 447-489.

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BALL, G.E. & ANDERSON, J.N. (1962) The taxonomy and speciation of Pseudophonus. The CatholicUniversity of America Press, Washington, D.C. 99 pp.

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BALL, G.E. & HILCHIE, G. J. (1983) Cymindine Lebiini of authors: redefinition and reclassificationof genera (Coleoptera: Carabidae). Quaestiones Entomologicae, 19, 93-216.

BALL, G.E. & MADDISON, D.R. (1987) Classification and evolutionary aspects of the species ofthe 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 withnotes about the speices in southwestern United States and selected species from Mexico.Quaestiones entomologicae, 26, 30-116.

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BALL, G.E. & NIMMO, A.P. (1983) Synopsis of the species of Subgenus Progaleritina Jeannel,including reconstructed phylogeny and geographical history (Coleoptera: Carabidae: GaleritaFabricius). Transactions of the American Entomological Society, 109, 295-356.

BALL, G.E. & ROUGHLEY, R. (1982) The Hypherpes-like taxa of southern Mexico: classificationand evolutionary considerations (Coleoptera: Carabidae: Pterostichus). Transactions of theAmerican Entomological Society, 108: 315-399.

BALL, G.E. & SHPELEY D. (1983) The species of eucheiloid Pericalina: Classification and evolutionaryconsiderations (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.

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CHAUDOIR, M. de (1835) Description de quelques genres et espèces de carabiques nouveaux.Annales de la Société Entomologique de France, 4, 429-448.

CHAUDOIR, M. de (1837a) Description de quelques genres nouveaux et de quelques espèces nouvellesou inédities de carabiques. Bulletin de la Société Impériale des Naturalistes de Moscou, 10(3), 3-20.

CHAUDOIR, M. de (1837b) Genres nouveaux et espèces nouvelles de coléoptères de la familledes carabiques. Bulletin de la Société Impériale des Naturalistes de Moscou, 10(7), 3-50.

CHAUDOIR, M. de (1838) Tableau d’une nouvelle subdivsion du genre Feronia Dejean suivi d’unecharactéristique de trois nouveaux genres de carabiques. Bulletin de la Société Impériale desNaturalistes de Moscou, 11, 3-32.

CHAUDOIR, M. de (1842a) Catalogue des Carabiques recueillis dans la province de Mazendéranprès d’Astrabat par Mr. de Karéline. Bulletin de la Société Impériale des Naturalistes de Moscou,15, 801-831.

CHAUDOIR, M. de (1842b) Description de quelques generes nouveaux de la famille des carabiques.Bulletin de la Société Impériale des Naturalistes de Moscou, 15, 832-857.

CHAUDOIR, M. de (1843a) Genres nouveaux de la famille des carabiques. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 16(III), 383-427.

CHAUDOIR, M. de (1843b) Carabiques nouveaux. Bulletin de la Société Impériale des Naturalistes deMoscou, 16, 671-791.

CHAUDOIR, M. de (1844) Trois mémoires sur la famille des carabiques. Bulletin de la Société Impérialedes Naturalistes de Moscou, 17, 415-479.

CHAUDOIR, M. de (1846a) Carabiques. Pp. 47-209. Carabiques nouveaux de la Crimée. Pp. 227-234. In, Chaudoir, M. de & Hochhuth, J.H., Enumération des Carabiques et Hydrocantharesrecueillis pendant un voyage au Caucase et dans les provinces transcaucasiennes par le Baron M. deChaudoir et le Baron A. de Gotsch. 268pp. J. Wallner, Kiew.

CHAUDOIR, M. de (1846b) Note sur le groupe des Stomides et description d’un nouveau genre decelui des Somoplatides. Bulletin de la Société Impériale des Naturalistes de Moscou, 19(2), 511-542.

CHAUDOIR, M. de (1847) Notes sur le genre Agra et desciption de plusieurs espèces nouvelles.Bulletin de la Société Impériale des Naturalistes de Moscou, 20, 87-113.

CHAUDOIR, M. de (1848) Mémoire sur la familla des carabiques. Première partie. Bulletin de laSociété Impériale des Naturalistes de Moscou, 21(1), 3-134.

CHAUDOIR, M. de (1850a) Mémoire sur la familla des carabiques. 2e partie. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 23(1), 3-85.

CHAUDOIR, M. de (1850b) Mémoire sur la familla des carabiques. 2e partie (continuation). Bulletinde la Société Impériale des Naturalistes de Moscou, 23(2), 349-460.

CHAUDOIR, M. de (1850c) Supplement à la faune des carabiques de la Russie. 4. Trois espècesnouvelles du groupe des Amaroides. Bulletin de la Société Impériale des Naturalistes de Moscou,23(3), 62-206.

CHAUDOIR, M. de (1852) Mémoire sur la familla des carabiques. 3e partie. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 25(1), 3-104.

CHAUDOIR, M. de (1854) Mémoire sur la familla des carabiques, 4e partie. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 27(1), 112-144, (2), 279-352.

CHAUDOIR, M. de (1855) Mémoire sur la familla des carabiques, 5e partie. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 28(1), 1-110.


CHAUDOIR, M. de (1856) Mémoire sur la familla des carabiques, 6e partie. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 29(3), 187-291.

CHAUDOIR, M. de (1857) Mémoire sur la familla des carabiques, 6e partie (continuation). Bulletinde la Société Impériale des Naturalistes de Moscou, 30(3), 1-64.

CHAUDOIR, M. de (1859a) Monographie du genre Colopodes MacLeay. Annales de la SociétéEntomologique de France, (ser. 3) 7, 287-364.

CHAUDOIR, M. de (1859b) Beitrag zur Kenntniss der europäischen Feroniden. EntomologischeZeitung, 20, 113-131.

CHAUDOIR, M. de (1860) Matériaux pour servir à l’étude des Cicindélètes et des Carabiques. 1e

partie. Bulletin de la Société Impériale des Naturalistes de Moscou, 33(4), 269-337.CHAUDOIR, M. de (1861a) Révision du Genre Agra. Annales de la Société Entomologique de France, 1,

109138.CHAUDOIR, M. de (1861b) Beitrag zur Kenntnis einger carabicien-Gattungen. Berliner Entomologische

Zeitschrift, 5, 116-131.CHAUDOIR, M. de (1861c) Matériaux pour server à l’étude des Cicindélètes et des Carabiques.

(Continuation). Bulletin de la Société Impériale des Naturalistes de Moscou, 34(2), 491-576.CHAUDOIR, M. de (1861d) Révision des espèces qui rentrent dans l’ancien genre Panagaeus. Bulletin

de la Société Impériale des Naturalistes de Moscou, 34(4), 335-360.CHAUDOIR, M. de (1862a) Description, sommaires d’espèces nouvelles de cicindélètes et de

carabiques de la collection de M. le Baron Chaudoir. Revue et Magasin de Zoologie Pure etAppliquée, (2), 14, 484490.

CHAUDOIR, M. de (1862b) Matériaux pour servir à l’étude des Cicindélètes et des Carabiques. 3e

partie. Bulletin de la Société Impériale des Naturalistes de Moscou, 35(4), 275-320.CHAUDOIR, M DE. (1863a) Description de Cicindélètes et de Carabiques nouveaux. Revue et

Magasin de Zoologie Pure et Appliquée, (2), 15, 111-120, 187-188, 223-225.CHAUDOIR, M. de (1863b) Enumération des Cicindélètes et de Carabiques et recuellis dans la Russie

méridionale, dans la Finlande septentrionale et dans la Sibérie orientale par MM. Alexandre etArtur de Nordmann. Bulletin de la Société Impériale des Naturalistes de Moscou, 36(1), 201-232.

CHAUDOIR, M. de (1863c) Description de quelque nouvelles espèces de Cicindélètes et deCarabiques. Annales de la Société Entomologique de France, 4(3), 447-450.

CHAUDOIR, M. de (1865) Catalogue de la collection de Cicindélétes de M. le Baron de Chaudoir, 64 pp. J.Nys, Bruxelles.

CHAUDOIR, M. de (1866a) Corrections et additions a la Revision du Genre Agra. Annales de laSociété Entomologique de France, 6, 77104.

CHAUDOIR, M. de (1866b) Supplément à la monographie du genre Pelecium (Beliner Entomol.Zeitschrift, 1865, p. 127). Revue et Magasin de Zoologie, 1866, 108-110.

CHAUDOIR, M. de (1867) Note sur le genre Oxystomus. Annales de la Société Entomologique de France,11, 133-135.

CHAUDOIR, M. de (1868a) Note monographique sur le Genre Omophron. Magasin de Zoologie Pureet Appliquée, (2), 20, 54-63.

CHAUDOIR, M. de (1868b) Observations synonymiques sur les Carabiques de l’Amériqueseptentrionale et description d’espèces nouvelles de ce pays. Revue et Magasin de Zoologie Pureet Appliquée, (2), 20, 161-171, 211-217, 239-245, 283301, 331-345.

CHAUDOIR, M. de (1868c) Révision du groupe des ozénides. Annales de la Société Entomologique deBelgique, 11, 43-74.

CHAUDOIR, M. de (1869a) Mémoire sur les coptodérides. Annales de la Société Entomologique deBelgique, 12, 163-256.


CHAUDOIR, M. de (1869b) Description de Cicindélètes et de Carabiques noveaux. Revue et Magasinde Zoologie Pure et Appliquée, (2), 21, 22-28, 64-70, 114-122, 170-173, 203-208.

CHAUDOIR, M. de (1869c) Descriptions de Calosoma nouveaux des collections de Mm. de Chaudoiret Sallé. Annales de la Société Entomologique de France, (4). 9, 367-378.

CHAUDOIR, M. de (1869d) Mémoire sur les thyréoptérides. Annales de la Société Entomologique deBelgique. 12, 113-162.

CHAUDOIR, M. de (1871a) Monographie des lébiides. 1e partie. Bulletin de la Société Impériale desnaturalistes de Moscou, 44, 1-87.

CHAUDOIR, M. de (1871b) Essai monographique sur le groupe des pogonides. Annales de laSociété Entomologique de Belgique, 14, 21-61.

CHAUDOIR, M. de (1871c) Monographie des lébiides. Bulletin de la Société Impériale des naturalistes deMoscou. 43(3)(1870), 111-255.

CHAUDOIR, M. de (1872a) Monographie des callidides. Annales de la Société Entomologique de Belgique,15, 97-204.

CHAUDOIR, M. de (1872b) Observations sur quelques genres de Carabiques avec la descriptiond’espèces nouvelles. Bulletin de la Société Impériale des naturalistes de Moscou, 45(2), 382-420.

CHAUDOIR, M. de (1872c) Descriptions d’espèces nouvelles de Carabiques de la tribu destroncatipennes, et remarques synonymiques. Revue et Magasin de Zoologie Pure et Appliquée,(2), 23, 101107, 138-143, 168-172, 212-221, 241-250.

CHAUDOIR, M. de (1873a) Essai monographique sur le genre Cymindis proprement dit. BerlinerEntomologische Zeitschrift, 17, 53-120.

CHAUDOIR, M. de (1873b) Matériaux pour servir à l’étude des féroniens. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 46(3), 85-116.

CHAUDOIR, M. de (1874) Matériaux pour servir à l’étude des féroniens. Suite. Bulletin de la SociétéImpériale des Naturalistes de Moscou, 48(1), 1-34.

CHAUDOIR, M. de (1875) Genres aberrants du groupe des cymindides. Bulletin de la Société Impérialedes Naturalistes de Moscou, 49(3), 1-61.

CHAUDOIR, M. de (1876a) Étude monographique des masoréides, des tetragonodérides et dugenre Nematotarsus. Bulletin de la Société Impériale des Naturalistes de Moscou, 51(3), 1-84.

CHAUDOIR, M. de (1876b) Notes et additions au memoire de M. Reed sur les carabiques du Chiliinsérè dans les Proceedings of the Zoological Society of Londan (January 1874: 48). Premièrepartie. Annales de la Société Entomologique de Belgique, 19, 105-124.

CHAUDOIR, M. de (1876c) Monographie de brachynides. Annales de la Société Entomologique deBelgique, 19, 11-104.

CHAUDOIR, M. de (1876d) Monographie du genre Poecilus. L’Abeille, Journal d’Entomologie, 14,1-54.

CHAUDOIR, M. de (1876e) Monographie des siagonides. Bulletin de la Société Impériale des Naturalistesde Moscou, 50(1), 62-125.

CHAUDOIR, M. de (1876f) Monographie des chléniens. Annali del Museo Civico di Storia Naturale diGenova, 8, 3-315.

CHAUDOIR, M. de (1876g) Catalogue des Cicindélètes et des Carabiques recueillis par M. AchileRaffray, en Abyssine, avec la description des espèces nouvelles. Revue et Magasin de ZoologiePure et Appliquée, (3), 4, 329-388.

CHAUDOIR, M. de (1877) Genres nouveaux et espèces inédites de la familla des Carabiques.Bulletin de la Société Impériale des Naturalistes de Moscou, 52(2), 188-268.

CHAUDOIR, M. de (1878a) Descriptions de genres nouveaux et d’ espèces inédites de la famillades carabiques. Bulletin de la Société Impériale des Naturalistes de Moscou, 53(3), 1-80.


CHAUDOIR, M. de (1878b) Révision des genres Onychopterygia, Dicranoncus et Colpodes. Annales de laSociété Entomologique de France, (5), 8, 275-336 [part].

CHAUDOIR, M. de (1879a) Monographie des scaritides (Scaritini). Première partie. Annales de laSociété Entomologique de Belgique, 22, 124-182.

CHAUDOIR, M. de (1879b) Révision des genres Onychopterygia, Dicranoncus et Colpodes. Annales de laSociété Entomologique de France, (5), 8, 337-382 [part].

CHAUDOIR, M. de (1879c) Essai monographique sur les Panagéides. Annales de la SociétéEntomologique de Belgique, 21(1878), 85-186.

CHAUDOIR, M. de (1880a) Monographie des scaritides (Scaritini). Deuxième partie. Annales de laSociété Entomologique de Belgique, 23, 5-130.

CHAUDOIR, M. de (1880b) (Descriptions de six nouvelles espèces de carabiques de genre Pasimachus.Bulletin de la Société Entomologique de France, 1880, lxxxv-lxxxviii.

CHAUDOIR, M. de (1880c) Essai monographique sur les morionides. Bulletin de la Société Impérialedes Naturalistes de Moscou, 55(2), 317-384.

CHAUDOIR, M. de (1882) Monographie des oodides, Première partie. Annales de la SociétéEntomologique de France, (6). 2, 317-378. [Ouvrage posthume]

CHAUDOIR, M. de (1883a) Monographie des oodides, Deuxième partie. Annales de la SociétéEntomologique de France, (6). 2, 485-554. [Ouvrage posthume].

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ERWIN, T.L. (1969a) A new bombardier beetle in Canada (Coleoptera: Carabidae). Opusculaentomologica, 34(3), 283286.

ERWIN, T.L. (1969b) Preliminary diagnoses of five new North American Brachinus (Coleoptera:Carabidae). Opuscula entomologica, 34(3), 287288.

ERWIN, T.L. (1970a) The nearctic species of the Genus Leistus Frolich (Coleoptera: Carabidae).PanPacific Entomologist, 46, 111119.

ERWIN, T.L. (1970b) A reclassification of bombardier beetles and a taxonomic revision of theNorth and Middle American species (Carabidae: Brachinida). Quaestiones entomologicae, 6,4215.

ERWIN, T.L. (1971a) Notes and corrections to a reclassification of bombardier beetles (Carabidae,Brachinida). Quaestiones entomologicae, 7, 281.

ERWIN, T.L. (1971b) Fossil tachyine beetles from Mexican and Baltic Amber with notes on anew synonymy of an extant group (Col., Carabidae). Entomologica Scandinavica 2(3), 233236.

ERWIN, T.L. (1972a) The lost type of Brachinus sallei Chaudoir (Coleoptera: Carabidae). PanPacificEntomologist, 48(1), 6364.

ERWIN, T.L. (1972b) Two new genera of bembidiine carabid beetles from Australia and SouthAmerica with notes on their phylogenetic and zoogeographic significance (Coleoptera).Breviora No. 383, 119.

ERWIN, T.L. (1973a) Studies of the subtribe Tachyina (Coleoptera: Carabidae: Bembidiini), PartI: A revision of the Neotropical genus Xystosomus Schaum. Smithsonian Contributions to Zoology,No. 140, 1-39.

ERWIN, T.L. (1973b) A supplement to the bombardier beetles of North and Middle America: Newrecords for Middle America (Coleoptera: Carabidae). The Coleopterists Bulletin, 27(2), 7982.

ERWIN, T.L. (1974a) Studies of the subtribe Tachyina (Coleoptera: Carabidae: Bembidiini), PartII: A revision of the New WorldAustralian Genus Pericompsus LeConte. SmithsonianContributions to Zoology, No. 162, 1-96.

ERWIN, T.L. (1974b) Studies of the subtribe Tachyina (Coleoptera: Carabidae: Bembidiini)Supplement A: Lectotype designations for New World species, two new genera, and noteson generic concepts. Proceedings of the Entomological Society of Washington, 76(2), 123155.

ERWIN, T.L. (1975) Studies of the subtribe Tachyina (Coleoptera: Carabidae:Bembidiini), PartIII: Systematics, phylogeny, and zoogeography of the GenusTachyta Kirby. SmithsonianContributions to Zoology, No. 208, 1-68.

ERWIN, T.L. (1978a) Studies of the subtribe Tachyina (Coleoptera: Carabidae: Bembidiini),Supplement C: Discovery of the sister group of Mioptachys Bates, in the Amazon BasinInpa psydroides a new genus and species. The Coleopterists Bulletin, 32(1), 2936.

ERWIN, T.L. (1978b) Systematic, natural history, and zoogeographic notes on the Genus AgraFabricus, with a description of a new species from Panamá (Coleoptera: Carabidae: Lebiini).The Coleopterists Bulletin, 32(4), 261268.

ERWIN, T.L. (1978c) The larva of Neotropical Enceladus gigas Bonelli (Coleoptera: Carabidae:Siagoninae: Enceladini) with notes on the phylogeny and classification of some of themore primitive tribes of ground beetles. The Coleopterists Bulletin, 32(2), 99106.

ERWIN, T.L. (1980) Systematic and zoogeographic notes on Apotomini, an old world faunalelement new to the western hemisphere list, with a description of a new species fromBrazil (Coleoptera: Carabidae). The Coleopterists Bulletin, 34(1), 99104.

ERWIN, T.L. (1981) Natural history of Plummers Island, Maryland. XXVI. The ground beetles of atemperate forest site (Coleoptera: Carabidae): An analysis of fauna in relation to size, habitatselection, vagility, seasonality, and extinction. Bulletin of the Biological Society of Washington, 5, 105224.


ERWIN, T.L. (1982a) Small terrestrial groundbeetles of Central America (Carabidae: Bembidiinaand Anillina). Proceedings of the California Academy of Sciences, 42(19), 455496.

ERWIN, T.L. (1982b) Agra, arboreal beetles of Neotropical forests: erythropus group systematics(Carabidae). Systematic Entomology, 7, 3971.

ERWIN, T.L. (1982c) Agra, arboreal beetles of Neotropical forests: platyscelis group systematics(Carabidae). Systematic Entomology, 7, 185210.

ERWIN, T.L. (1983) Agra, arboreal beetles of Neotropical forests: famula and formicaria groupsystematics (Carabidae). Systematic Entomology, 8, 263292.

ERWIN, T.L. (1984a) Agra, arboreal beetles of Neotropical forests: palmata group systematics(Carabidae). Systematic Entomology, 9, 948.

ERWIN, T.L. (1984b) Small terrestrial ground beetles of the Amazon Basin (Coleoptera: Bembidiini:Tachyina and Anillina). Amazoniana, 8(4), 511518.

ERWIN, T.L. (1985) The taxon pulse: A general pattern of lineage radiation and extinction amongcarabid beetles, pp. 437-472. In, Ball, G. E. (ed.). Taxonomy, phylogeny, and zoogeographyof beetles and ants: A volume dedicated to the memory of Philip Jackson Darlington Jr.1904 -1983. Dr. W. Junk b.v. Publishers, The Hague.

ERWIN, T.L. (1986) Agra, arboreal beetles of Neotropical forests: mixtagroup, virgatagroup, andohausigroup systematics (Carabidae). Systematic Entomology, 11, 293316.

ERWIN, T. L. (1987) Agra, arboreal beetles of Neotropical forests: feisthameli group systematics(Carabidae). Systematic Entomology, 12, 137161.

Erwin, T.L. (1991). Natural history of the carabid beetles at the BIOLAT Rio Manu BiologicalStation, Pakitza, Perú. Revista Peruana de Entomologia, (1990) 33:1-85.

ERWIN, T.L. (1993) Agra, arboreal beetles of Neotropical forests: rufoaenea and quararibea groupsystematics (Carabidae). Revista peruana de Entomologia, 8, 263292.

ERWIN, T.L. (1994) Arboreal beetles of tropical forests: The Xystosomi group, SubtribeXystosomina (Coleoptera: Carabidae: Bembidiini). Part I. Character analysis, taxonomy,and distribution. The Canadian Entomologist, 126(3), 549-666.

ERWIN, T.L. (2000a) Arboreal beetles of Neotropical forests: Agra Fabricius, the Novaurora andDimidiata Groups (Coleoptera: Carabidae: Lebiini: Calleidina).Smithsonian Contributions toZoology, No. 608, pp. i-iv, 1-33.

ERWIN, T.L. (2000b) Arboreal beetles of Neotropical Forests: Taxonomic supplement for theplatyscelis group with new species and distribution records (Coleoptera: Carabidae, Lebiini,Agrina). The Coleopterists Bulletin, 54(1), 90-119.

ERWIN, T.L. (2000c) Arboreal beetles of Neotropical Forests: Taxonomic supplement for theAgra virgata and ohausi groups with a new species and additional distribution records(Coleoptera: Carabidae). The Coleopterists Bulletin, 54(2), 251-262.

ERWIN, T.L. (2000d) A new genus and species of Lachnophorini and two newspecies of Lebiinifrom Costa Rica (Coleoptera: Carabidae). TheColeopterists Bulletin, 54(3), 279-283.

ERWIN, T.L. (2002) The Beetle Family Carabidae of Costa Rica: Twenty-nine new species ofCosta Rican Agra Fabricius 1801 (Coleoptera: Carabidae, Lebiini, Agrina). Zootaxa, 119, 1-68.

ERWIN, T.L. (2004a) The Beetle Family Carabidae of Costa Rica and Panamá: Descriptions offour new genera and six new species with notes on their way of Life (Insecta: Coleoptera).Zootaxa, 537, 1-18.

ERWIN, T.L. (2004b) The Beetle Family Carabidae of Costa Rica: The Genera of the CryptobatidaGroup of Subtribe Agrina, Tribe Lebiini, with new species and notes on their way of life(Insecta: Coleoptera). Zootaxa, 662, 1-54.


ERWIN, T.L. (2004c) The beetle family Carabidae of Costa Rica: The genus Epikastea Liebke ofthe Plochonida Group, with new Neotropical species and notes on their way of life (Insecta:Coleoptera, Lebiini, Agrina). Zootaxa, 790, 1-20.

ERWIN, T.L. & BALL, G.E. (1972) Classification of the Ovipennis and Trifaria groups of NebriaLatreille (Coleoptera: Carabidae: Nebriini). Proceedings of the Biological Society of Washington,85(7), 77108.

ERWIN, T.L. & KAVANAUGH, D.H. (1981) Systematics and zoogeography of Bembidion Latreille: I.The carlhi and erasum groups of western North America (Coleoptera: Carabidae, Bembidiini).Entomologica Scandinavica, Supplement 15, 33-72.

ERWIN, T.L. & STORK, N. (1985) The Hiletini, an ancient and enigmatic tribe of Carabidae witha pantropical distribution (Coleoptera). Systematic Entomology, 10, 405451.

ERWIN, T.L., & KAVANAUGH, D.H. (1999) Studies of the Subtribe Tachyina (Coleoptera:Carabidae:Bembidiini), Supplement D: Description of a Male of Costitachys inusitatus Erwin, withNotes on Distribution of this Species. The ColeopteristsBulletin, 53(1), 52-55.

ERWIN, T.L. & KAVANAUGH, D.H. (2000) A New Species of the Tachyta SubgenusParatachytaErwin (Coleoptera: Carabidae: Bembidiini: Tachyina) fromSoutheastern China with a NewKey and Analysis of PhylogeneticRelationships among Species of the Subgenus, Pp. 183-194. In, Zamotajlov, A. & Sciaky R. (eds.). Advances in carabidology (Papers dedicated to thememory of Prof. Oleg L. Kryzhanovskij). MUISOpublications, Krasnodar, 473pp.

ERWIN, T.L. & MEDINA, R.F. (2003) The Neotropical species Askalaphium depressum (Bates):Larval description, first diagnosis and illustrations of immature Ctenodactylini, with naturalhistory notes on the genus and tribe (Coleoptera: Carabidae). Zootaxa, 273, 1-15.

ERWIN, T.L. & ASCHERO, V. (2004b) Cicindis horni Bruch (Coleoptera: Carabidae, Cicindini): The FairyShrimp Hunting Beetle, its way of life on the Salinas Grandes of Argentina. Zootaxa, 553, 1-16.

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ETONTI, M. & MATEU, J. (1992a) Sette nouvi Trechinae del Perú. Bollettino del Museo Civico di StoriaNaturale de Venecia, 41 (1990), 109-130.

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Leyden Museum, 17, 15-16.HORN, W. (1895d) Matériaux pour servir à l’étude des cicindélides. Anales el Museo Nacional de

Historia Natural de Buenos Aires, 4, 173-176.HORN, W. (1896a) Quatre espèces nouvelles de coléoptères (famille des cicindélides) du Muséum

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Zeitschrift, 1898, 193-198.HORN, W. (1898c) Ten new species of Cicindelidae. Notes from the Leyden Museum, 20, 101-108.HORN, W. (1900a) 7 neue Cicindeleten. Deutsche Entomologische Zeitschrift, 1900, 361-365.HORN, W. (1900b) De novis Cicindelidarum speciebus. Deutsche Entomologische Zeitschrift, 1900,

193-212.HORN, W. (1900c) De duabus novis generic tetrachae speciebus ex Ecuadoria. Entomologisches

Nachrichtenblatt, 26, 53-54.HORN, W. (1900d) Über einiger U.S.A. – Cicindelen. Entomologisches Nachrichtenblatt, 26(8), 116-119.HORN, W. (1901a) De 7 novis Cicindelidis. Deutsche Entomologische Zeitschrift, 1901, 353-358.HORN, W. (1901b) Über Oxygonia floridula Bat. und gloriola Bat. Deutsche Entomologische Zeitschrift,

1901, 124.HORN, W. (1902) Briefe eines reisenden Entomologen. Deutsche Entomologische Zeitschrift, 1902,

369-396.HORN, W. (1903b) Briefe eines reisenden Entomologen. Deutsche Entomologische Zeitschrift, (1903)

177-198.HORN, W. (1903c) List of the Cicindelidae of Mexico and on their relationship with the species

of the United States. Journal of the New York Entomological Society, 11, 213-221.HORN, W. (1903d) On the Cicindelidae of southern Venezuela, collected by Edward A. Klages,

1898-1900. Proceeding of the Entomological Society of Washington, 5, 331-334.HORN, W. (1904) Über die Cicindeliden-Sammlungen von Paris und London. Deutsche

Entomologische Zeitschrift, 1904, 81-99.HORN, W. (1905a) Systematischer Index der Cicindeliden, Deutsche Entomologische Zeitschrift, Beiheft,

1, 1-56.HORN, W. (1905b) Two new species of Cicindelidae. Notes from the Leyden Museum, 25, 219-220.HORN, W. (1905c) Die Gattungen Oxygonia und Odontochila. Deutsche Entomologische Zeitschrift,

1905, 145-149.HORN, W. (1906a) Über die neotropishen Arten der Cicindela argentata-Gruppe. Deutsche

Entomologische Zeitschrift, 1906, 87-92.HORN, W. (1906b) Odontochila lacordairei Gory rhytidopteroides (nov. Subsp.) und über das Vorkommen

verschiedener Rassen derselben Species am selben Ort. Deutsche Entomologische Zeitschrift,1906, 175-176.

HORN, W. (1907a) Zur Kenntnis der Gattung Cicindela. Deutsche Entomologische Zeitschrift, 1907, 20-25.HORN, W. (1907b) Neues über Odontochila, Cicindela, und Dromica. Stettiner Entomologische Zeitung,

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186-188.HORN, W. (1909b) Zwei neue neotropische Megacephalae (Col.). Deutsche Entomologische Zeitschrift,

1909, 284-286.HORN, W. (1909c) Contribution a l’étude des Cicindelinae (Col.). Annales de la Société Entomologique

de Belgique, 53, 443-447.HORN, W. (1910a) Family Carabidae: Subfamily Cicindelinae. Genera Insectorum, 82B, 105-208,

plates 6-15.HORN, W. (1910b) Une forme nouvelle d’Omus (Cicindel. Col.). Annales de la Société Entomologique

de Belgique, 54, 293-295.HORN, W. (1911) Drei neue südamerikanishche Ctenostoma-Arten. Entomologische Blätter, 8, 152-155.HORN, W. (1913a) 50 neue Cicindelinae. Archiv für Naturgeschichte, (A) 78, 1-33. Berlin.HORN, W. (1913b) Diesjaahrige Omus-Funde von F.W. Nunenmacher (Col.). Entomologische

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MANDL, K. (1955) Ergebnisse einer Revision der Carabiden-Sammlung des Natur-historischenMuseums Wien. (4. Teil). Annalen des Naturhistorischen Museums in Wien, 60(1954-1955), 236-272.

MANDL, K. (1956a) Neun neue Cicindelidenformen aus tropischen Ländern. 2. Zur Kenntnisder Cicindeliden-Fauna von Bolivien. Entomologischen Arbeiten aus dem Muséum G. Frey, Tutzing,7, 383-390.

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Rundschau, 39, 24-25.MANDL, K. (1963a) Neue Cicindelidae-Formen aus tropischen Ländern. Koleopterologische

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(mit 2 Tafeln). Entomologischen Arbeiten aus dem Muséum G. Frey, Tutzing, 14, 575-592.MANDL, K. (1964) Ergebnisse einer Forschungsreise in das Rio Negro – (Amazonas) Gebiet in

Brasilien. Beschreibung einer neuer Megacephala – (Phaeoxantha–) Art. KoleopterologischeRundschau, 42, 15-17.

MANDL, K. (1967a) Zweiter Beiträg zur Kenntnis der Cicindelidae-Formen von Peru (Col.).Entomologischen Arbeiten aus dem Muséum G. Frey, Tutzing, 17, 432-439.

MANDL, K. (1967b) Neue Cicindeliden – Formen in der Sammlung des museums G. Frey,Tutzing (Col.). Entomologischen Arbeiten aus dem Muséum G. Frey, Tutzing, 18, 384-387.

MANDL, K. (1970) Neue Cicindelidae-Formen aus der Sammlung des zoologischen Museumsder Humboldt-Universität zu Berlin. Zeitschrift der Arbeitsgemeinschaft Österreichischer entomologen,22, 65-80.

MANDL, K. (1971) Über neue und wenig bekannte Formen der Familie Carabidae, 2. Zeitschriftder Arbeitsgemeinschaft Österreichischer entomologen, 23, 40-48.

MANDL, K. (1972)MANDL, K. (1973a) Neue Cicindelidae-Formen aus den tropischen Gebieten Afrikas und Süd-

Amerikas. A Fünf neue Brasiella-Formen aus Süd-Amerika. Entomologischen Arbeiten aus demMuséum G. Frey, Tutzing, 24, 290-303.

MANDL, K. (1973b) Fragmenta carabologica. Beschreibung einer neuen Untergattung und Zweierneuer Rassen. Mitteilungen Entomologischen Gesellschaft, Basel, (N.F.) 23, 77-89.

MANDL, K. (1975a) Neue Cicindelidae – Formen aus Südamerika (Coleoptera). Zeitschrift derArbeitsgemeinschaft Österreichischer entomologen, 1974 (1975), 26(1), 15-22.

MANDL, K. (1975b) Weitere Beiträge zur Kenntnis der Carabini (Carabidae, Col.). KoleopterologischeRundschau, 52, 61-85.

MANDL, K. (1977) Ceroglossus chilensis nigritulus, eine neue subspecies aus Argentinien (Col.Carabidae). Mitteilungen Entomologischen Gesellschaft, Basel, 27(1), 6-7.

MANDL, K. (1981a) Eine Cicindeliden-Ausbeute aus dem nördlichen Brasilien und Beschreibungneuer Arten aus dieser Familie (Col., Cicindelidae). Entomologica Basiliensia, 6, 154-160.

MANDL, K. (1981b) Neun neue Formen aus der Familie Cicindelidae aus fünf Kontinenten(Col.). Koleopterologische Rundschau, 55, 3-18.


MANDL, K. (1983) Cicindela (Brasiella) dominicana, eine neue Art aus West-Indien (Cicindelidae,Col.). Zeitschrift der Arbeitsgemeinschaft Oesterreichischer Entomologen, 34(3-4), 117-119.

MANDL, K. (1992) Ein Beitrag zur Kenntnis einiger Chlaeniinae-(Callistinae-) Formen aus derpalaarktischen und der orientalischen Region. (Zugleich zweite Auswertung der vomNaturhistorischen Museum in Wien erworbenen Chlaeniinae-Spezialsammlung ErnstGrundmanns). (Carabidae: Coleoptera). 2. Teil. Annalen des Naturhistorischen Museums in WienSerie B Botanik und Zoologie, 93, 59-103.

MANEE, A.H. (1915) Three new species of Coleoptera from North Carolina. Entomological News,26, 175-176.

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MANNERHEIM, C.G. VON (1843) Beitrag zur Käferfauna des Aleutischen Inseln, der Insel Sitkaund Neu Californiens. Bulletin de la Société Impériale des Naturalistes de Moscou, 16(2), 175-324

MANNERHEIM, C.G. VON (1844) Lettre a S. E. Mr. Fischer de Waldheim ou relation d’un voyagefait en 1844, en Denmark et dans le nord de l’Allemagne. Bulletin de la Société Impériale desNaturalistes de Moscou, 17, 844-872.

MANNERHEIM, C.G. VON (1852) Zweiter Nachtrag zur Käfer-Fauna der nord-amerikanischen Laenderdes russischen Reiches. Bulletin de la Société Impériale des Naturalistes de Moscou, 25(1), 283-387.

MANNERHEIM, C.G. VON (1853) Beitrag zur Käferfauna des Aleutischen Inseln, der Insel Sitkaund Neu Californiens. Bulletin de la Société Impériale des Naturalistes de Moscou, 26(2), 95-273.

MAŘAN, J. (1934) De novis carabidarum speciebus formisque ex insula Creta. SborníkEntomologického Oddeliní Národního Musea v Praze, 12, 125-128.

MARES, A. (1921) A new species and a new variety of Cicndela (Col.). Entomological News, 32(10), 310.MARSHAM, T. (1802) Entomologia Britaninica, sistens insecta Britanniae indigena, secundum methodum

Linnaeanum disposita. Tomus I. Coleoptera. xxxi + 548pp. White, London.MARTORELL, L.F. (1939) Insects observed in the State of Aragua, Venezuela, South America.

Journal of Agriculture of the University of Puerto Rico. 23, 177-262.MATEU, J. (1953a) Algunos Tachyura del sudeste de España. Estratto dal Bollettino della Societa

Entomologica Italiana, 82(9-10), 95-98.MATEU, J. (1953b) Notas sobre carábidos españoles. (1.a note). Archivos del Instituto de Aclimatación,

Almeria, 1, 135-142.MATEU, J. (1954a) El genero Callidula Chaudoir. Revista de la Sociedad Entomológica Argentina, 17, 3-8.MATEU, J. (1954b) Notas sobre carabidos españoles (2ª nota). Bulletin de l’Institut Royal des Sciences

Naturelles de Belgique, 30(31), 1-8.MATEU, J. (1955) Revisión de los géneros Mimodromius Chaudoir y Mimodromites Mateu (Col.

Carabidae). Archivos del Instituto de Aclimatación. Almeria, 4 (1955), 63-108.MATEU, J. (1956) El genero Cymindidius Chaudoir (Col. Carabidae). Anales de Parques Nacionales,

6, 15-24.MATEU, J. (1959a) Nuevos datos sobre los Mimodromius Chaudoir (Coleoptera, Carabidae). Annali

del Museo Civico di Storia Naturale “G. Doria,” 3, no. 103, 1-11.MATEU, J. (1959b) El genero Cymindidius Chaudoir (Coleopteros, Carabidos). Bulletin de l’Institut

Royal des Sciences Naturelles de Belgique, 35(26), 1-8.MATEU, J. (1960) Cuarta note sobre el genero Mimodromius Chaudoir. Revista de la Sociedad Uruguaya

Entmolología, 4, 29-41.


MATEU, J. (1961) Sexta nota sobre los Lebiidae neotropicales (Coleópteros – Carábidos). Annalidel Museo Civico di Storia Naturale di Genova, 72, 161-178.

MATEU, J. (1964) Description de un nuevo Mimodromius Chaudoir de Chile y comentarios sobrediversas otras especies del genero (Coleoptera- Lebiidae). Revista de la Sociedad UruguayaEntmolología, 6, 7-12.

MATEU, J. (1968) Sobre algunos Dromiini (Col. Carabidae) nuevos o poco conocidos de AmericaNeotropical. Revista de la Sociedad Uruguaya Entmolología, 7, 72-79.

MATEU, J. (1970a) Notas sobre el género Monnea nov. De la Argentina y del Uruguay. Arquivosdo Museu Bocage, Series 2, 11, 139-148.

MATEU, J. (1970b) Sur un nouveau Mimodromius des montagnes de l’Écuador, pp. 173-179. In,Leleup, N. & Leleup, J., Mission zooogique belge aux î les Galápagos et en Ecuador. Vol. 2. Tervuren.

MATEU, J. (1970c) La répartition transandine du genre Mimodromius Chaudoir et descriptiond’une nouvelle espèce de l’Équateur. Bulletin de la Bulletin de la Société Entomologique de France,75, 150-156.

MATEU, J. (1970d) Sobre algunos Mimodromius chilenos y peruanos. Miscellanea Zoologia, 2(5), 77-91.MATEU, J. (1972a) Nouveaux Carabiques neotropicaux. Entomologischen Arbeiten aus dem Muséum

G. Frey, Tutzing, 1972, 321-330.MATEU, J. (1972b) Révision du genre Trechisibus Motsch. et genres voisins. Nouvelle Revue

d’Entomologie, 2, 53-72.MATEU, J. (1973) Los Dromius Bonelli de Améique. Anales de la Escuela Nacional de Ciencias Biológicas,

20, 89-121.MATEU, J. (1974a) Un nuevo Amblychila Say, procedente del altiplano mexicano (Coleopt.

Cicindelidae). Anales de la Escuela Nacional de Ciencias Biológicas, 21, 145-153.MATEU, J. (1974b) Sur les Microlestes Schmidt-Goebel du Méxique (Col. Carabidae Lebiinae).

Entomologischen Arbeiten aus dem Muséum G. Frey, Tutzing, 25, 261-275.MATEU, J. (1974c) Sobre algunos linajes de carabidos boreo-montanos de Mexico y sus relaciones

con el poblamiento entomologico del sistema volcanico transversal. Revista de la SociedadMexican de Historia Natural, 35, 181-224.

MATEU, J. (1975) Neuvas aportaciones al conocimiento del género Negrea Mateu y a su reparticion enel contiente americano (Coleoptera, Carabidae). Revista Brasileira de Entomologia, 19(2), 63-72.

MATEU, J. (1976a) Un notable endemismo de las montañas de México: El genero XenodromiusBates. Boletím da Sociedade portuguese do Ciéncias natural, 16, 137-147.

MATEU, J. (1976b) Sobre un nuevo Pterostichinae de Venezuela (Coleoptera, Carabidae). RevistaBrasileira de Entomologia, 20(2), 67-70.

MATEU, J. (1976c) Sobre dos nuevos géneros de Calleidini (Coleopt. Carabidae) procedentes deChile, Apterodromites nov. y Falsodromius nov. Miscellanea Zoologia, 3, 97-108.

MATEU, J. (1978a) Màs datos sobre los Trechisibus del Perú. Boletín de Entomología Venezolana (N.S.),1(3), 25-36.

MATEU, J. (1978b) Speleodesmoides raveloi, nuevo género y especie de carabido troglobio en unacueva de Venezuela. Boletín de Sociedad Venezolana Espeleologia (N.S.), 9(17), 21-28.

MATEU, J. (1979a) Màs datos sobre los Trechisibus del Perú (Nota 2). Miscellanea Zoologia, 5, 47-58.MATEU, J. (1979b) Nouvelles donnée sur les Dromius Bonelli de l‘Amerique Néotropicale. Nouvelle

Revue d’Entomologie, 9, 11-16.MATEU, J. (1979c) Deux nouveaux Agonini du Mexique (Col. Carabidae). Bulletin de la Société

Entomologique de France, 83 (1978), 182-186.MATEU, J. (1980) Un nouveau Anillini tetramère, gen. nov. & n. sp., recolté dans les grottes du

Pérou. Mémoires Biospéologique, 7, 201-207.


MATEU, J. (1981a) Révision de los Zuphium Latreille del continete Americano (Coleoptera:Carabidae). Folia Entomologica Mexicana, 47, 111-128.

MATEU, J. (1981b) Nuevos datos sobre la serie filetica de los Paratrechus Jeannel (Coleoptera,Carabidae). Descripcion de un nuevo genero de Venezuela y de una especie inedita deCosta Rica. Boletin de Entomologia Venezolana, 2(1), 1-10.

MATEU, J. (1982a) Una nueva especie del género Dromius Bonelli procedente de Colombia.Boletín de Entomología Venezolana (N.S.), 2(8), 69-72.

MATEU, J. (1982b) Columbitrechus g. nov., de la serie filética de los Paratrechus Jeannel y un nuevoOxytrechus de los Andes de Colombia. Eos, 58, 203-216.

MATEU, J. (1982c) Un interesante endemismo neotropical El Género Carbonellia Mateu (ColeopteraCarabidae Lebiinae). Boletín de Entomología Venezolana (N.S.), 2(5), 45-56.

MATEU, J. (1982d) Chaudoirella reichardt nouveau genre et n. sp. de Patriziini de l’Amériquenéotropicale. Nouvelle Revue d’Entomologie, 12(1), 45-51.

MATEU, J. (1982e) Le genre Negrea Mateu et sa distribution Néotropicale (1) (Coleoptera CarabidaeLebiinae). Annales de la Société Entomologique de France, N.S., 18(1), 89-105.

MATEU, J. (1983) Gen. Pseudomonnea nov., nouveau genre de Lebiinae de l’Amérique meridionale.Nouvelle Revue d’Entomologie, 13(3), 281-289.

MATEU, J. (1984a) Un nuevo subgénero y una especie inédita del género Mimodromius Chaudoir (ColeopteraCarabidae Lebiinae). Estratto dagli annali del Museo Civico di Storia Naturale di Genova, 85, 51-65.

MATEU, J. (1984b) Notas sobre una nueva especie del genero Prostolonis Mateu (Col., Carabidae).Acta Zoologica Lilloana, 37(2), 183-186.

MATEU, J. (1985) Reichardtina nom. nov. pour Reichardtia Mateu, 1982 (Col. Carabidae). NouvelleRevue d’Entomologie, 2(3), 266.

MATEU, J. (1987) Los Mimodromius del subgenero Gutierrezia Mateu (Coleoptera Carabidae Lebiinae).Annali-del-Museo-Civico-di-Storia-Naturale-”Giacomo-Doria,” 86 (1986), 217-224.

MATEU, J. (1988) Nouvelles espèces du genre Oxytrechus Jeannel recoltées en Équateur (Coleoptera,Carabidae). Nouvelle Revue d’Entomologie, 5(4), 305-313.

MATEU, J. (1989) Nuevos datos sobre los eucheiloidos sudamericanos (Coleoptera, Carabidae,Lebiinae). Elytron, 3, 61-67.

MATEU, J. (1991a) Trois nouvelles espèces du genre Dromius Bonelli (Coleoptera, Carabidae).Nouvelle Revue d’Entomologie, 8(4), 377-384.

MATEU, J. (1991b) Dromius martae n. sp., de la Colombia. Bulletin de la Société Entomologique deFrance, 96(1), 87-91.

MATEU, J. (1991c) Sur le genre Oxytrechus Jeannel, 1927, avec la description de nouvelles especesde l’Equateur et de la Colombie (Coleoptera, Carabidae, Trechinae). Eos, 67, 71-83

MATEU, J. (1993a) Sur deux nouveaux Carabiques Lebiinae de l’Amerique du Sud (Coleoptera,Carabidae). Elytron, 7, 157-164.

MATEU, J. (1993b) Mimodromius onorei nov. sp. del Ecuador y nuevos datos sobre otras especiesdel género (Coleoptera, Carabidae). Nouvelle Revue d’Entomologie, 10(4), 349-357.

MATEU, J. (1995) Nuevas especies de Pentagonicini y de Lebiinae de los Estados Unidos deAmérica (Coleoptera, Carabidae). Miscellania Zoologica (Barcelona), 17 (1993), 141-148.

MATEU, J. (1999) Contribution à la connaissance du genre Paratrechus Jeannel (Coleoptera,Carabidae, Trechini). Nouvelle Revue d’Entomologie, (1998) 15(4), 371-390.

MATEU, J. (2000a) Dos nuevos carabidos sudamericanos pertenecientes a las tribus Trechini yPterostichini (Coleoptera, Carabidae). Nouvelle Revue d’Entomologie, 17(1), 5-9.

MATEU, J. (2000b) Nuevas especies del genero Polpochila Solier, procedentes de America meridional(Coleoptera, Carabidae). Sessio Conjunta d’Entomologia ICHN SCL, 10, 117-122.


MATEU, J. (2003) Escolatrechus bolivianus gen. n., sp. n. de Trechinae de l’haut Plateau Bolivien(Coleoptera: Carabidae: Trechinae). Elytron (Barcelona), 16, 23-28.

MATEU, J. & BELLÉS, X. (1981a) Nuevos Trechidae del Perú: El genero Andinorites nov. Eos(1979-1980), 55-56, 131-142.

MATEU, J. & BELLÉS, X. (1981b) Résultats faunistiques de diverses campagnes réalisées enAmérique latine espèces inédites de Trechisibus Motschulsky récoltées au Pérou. NouvelleRevue d’Entomologie, 11(1), 39-45.

MATEU, J. & BELLÉS, X. (1982) Dos nuevos generos de Trechinae del Perú. Folia EntomologicaMexicana, 53, 67-74.

MATEU, J. & ETONI, M. (2002a) Due nuove specie peruviane del genere Trechisibus Motsch, 1863(Coleoptera, Carabidae, Trechinae). Atti del Museo Civico de Storia Naturale di Trieste, 49, 113-119.

MATEU, J. & ETONI, M. (2002b) Perucharidius andinus gen. n., sp. n. de anillini del Peru settentrionale(Coleoptera, Carabidae, Anillini). Atti del Museo Civico de Storia Naturale di Trieste, 49, 129-132.

MATEU, J. & ETONI, M. (2006) Due nuovi trechini di Cajamarca (Perù) (Coleoptera, Carabidae).Bolletino del Museo Civico di Storia Naturale di Verona, Botanica Zoología, 30, 147-150.

MATEU, J. & MORET, P. (2001) Cinq nouveaux Paratrechus de l’Ecuateur (Coleoptera, Carabidae,Trechini). Revue Française d’Entomologie, [N.S.] 23(1), 93100.

MATEU, J. & NÈGRE, J. (1972) Révision du genre Trechisibus Motsch. et genres voisins. NouvelleRevue d’Entomologie, 2(1), 53-72.

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INDEX | 317

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

Index

Calosoma (Blaptosoma) porosifrons Bates 101Calosoma (Blaptosoma) viridisulcatum laetulum (Jeannel) 107Calosoma (Blaptosoma) viridisulcatum viridilucens

Beheim & Breuning 107Calosoma (Blaptosoma) viridisulcatum viridisulcatu

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


Calosoma (Carabosoma) striatipenne Chaudoir 105Calosoma (Castrida) abbreviatum Chaudoir 85Calosoma (Castrida) alternans (Fabricius) 86Calosoma (Castrida) argentinensis Csiki 87Calosoma (Castrida) fulgens Chaudoir 93Calosoma (Castrida) galapageium Hope 93Calosoma (Castrida) granatense darwinia Van Dyke 94Calosoma (Castrida) granatense floreanum (Basilewsky) 94Calosoma (Castrida) granatense granatense Géhin 95Calosoma (Castrida) leleuporum (Basilewsky) 97Calosoma (Castrida) linelli Mutchler 98Calosoma (Castrida) peruviense Mandl 101Calosoma (Castrida) retusum (Fabricius) 102Calosoma (Castrida) rufipenne Dejean 103Calosoma (Castrida) sayi Dejean 103Calosoma (Castrida) trapezipenne Chaudoir 106Calosoma (Castrida) vagans Dejean 106Carabina Latreille 73Carabini Latreille 73Carabus Linné 108Carabus (Archicarabus) nemoralis nemoralis Müller 114Carabus (Aulonocarabus) truncaticollis truncaticol

Eschscholtz 116Carabus (Carabus) goryi Dejean 111Carabus (Carabus) granulatus granulatus Linné 111Carabus (Carabus) granulatus hibernicus Lindroth 112Carabus (Carabus) vinctus (Weber) 117Carabus (Diocarabus) chaimssonis Fischer von

Waldheim 109Carabus (Hemicarabus) serratus Say 114Carabus (Homoeocarabus) maeander maeander Fischer

von Waldheim 113Carabus (Megodontus) vietinghoffi vietinghoffi Adams 117Carabus (Oreocarabus) taedatus agassii LeConte 115Carabus (Oreocarabus) taedatus taedatus Fabricius 116Carabus (Tachypus) auratus Linné 108Carabus (Tachypus) cancellatus Illiger 109Carabus (Tanaocarabus) finitimus Haldeman 110Carabus (Tanaocarabus) forreri forreri Bates 110Carabus (Tanaocarabus) forreri willi Deuve 110Carabus (Tanaocarabus) hendrichsi Bolívar y Pieltain 113Carabus (Tanaocarabus) sylvosus Say 115Ceroglossina Lapouge 118Ceroglossus Solier 118Ceroglossus buqueti andestus Kraatz-Koschlau 118Ceroglossus buqueti argentinensis Jiroux 119Ceroglossus buqueti arieli Jiroux 119

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

INDEX | 319

Ceroglossus magellanicus ortrudheinzae Heinz & Jirzoux 134Ceroglossus magellanicus similis Kraatz-Koschlau 135Ceroglossus ochseni (Germain) 135Ceroglossus speciosus Gerstaecker 135Ceroglossus suturalis ancudanus Morawitz 136Ceroglossus suturalis olivaceus Kraatz-Koschlau 136Ceroglossus suturalis puxel Jiroux 136Ceroglossus suturalis suturalis (Fabricius) 137Cicindini Csiki 61Cicindis Bruch 62Cicindis horni Bruch 62Collyridini Brullé 171Ctenostoma Klug 172Ctenostoma (Ctenostoma) acciavatti Naviaux 173Ctenostoma (Ctenostoma) bahiaense Naviaux 175Ctenostoma (Ctenostoma) batesii Chaudoir 175Ctenostoma (Ctenostoma) chaudoiri (W. Horn) 178Ctenostoma (Ctenostoma) formicarium (Fabricius) 183Ctenostoma (Ctenostoma) guyanense W. Horn 185Ctenostoma (Ctenostoma) heydeni W. Horn 185Ctenostoma (Ctenostoma) jekelii jekelii Chevrolat 188Ctenostoma (Ctenostoma) jekelii raresculptum W. Horn 189Ctenostoma (Ctenostoma) rugicolle W. Horn 199Ctenostoma (Ctenostoma) rugiferum pseudoheydeni W. Horn 199Ctenostoma (Ctenostoma) rugiferum rugiferum (W. Horn) 199Ctenostoma (Ctenostoma) succinctum (Laporte de

Castlnau) 202Ctenostoma (Ctenostoma) transversum Naviaux 203Ctenostoma (Euctenostoma) bifasciatum bifasciatum

Dejean 176Ctenostoma (Euctenostoma) bifasciatum horni Naviaux 176Ctenostoma (Euctenostoma) bondari W. Horn 176Ctenostoma (Euctenostoma) eburatum Bates 182Ctenostoma (Euctenostoma) ferum Naviaux 183Ctenostoma (Euctenostoma) fryi Chaudoir 184Ctenostoma (Euctenostoma) inca Naviaux 187Ctenostoma (Euctenostoma) klugeanum W. Horn 189Ctenostoma (Euctenostoma) luctuosum Chaudoir 190Ctenostoma (Euctenostoma) magnum Naviaux 192Ctenostoma (Euctenostoma) monnei Naviaux 193Ctenostoma (Euctenostoma) rapillyi Naviaux 198Ctenostoma (Euctenostoma) regium Naviaux 198Ctenostoma (Euctenostoma) rugosum Klug 200Ctenostoma (Euctenostoma) sahlbergi Chaudoir 200Ctenostoma (Euctenostoma) sumlini Naviaux 203Ctenostoma (Euctenostoma) trinotatum (Fischer von

Waldheim) 203

Ctenostoma (Euctenostoma) tyrannum (Thomson) 204Ctenostoma (Euctenostoma) wiesneri Naviaux 206Ctenostoma (Microprocephalus) brevilabre W. Horn 177Ctenostoma (Microprocephalus) pusillum minimum Naviaux 197Ctenostoma (Microprocephalus) pusillum pusillum Naviaux 197Ctenostoma (Myrmecilla) abbreviatum Naviaux 172Ctenostoma (Myrmecilla) agnatum agnatum Chaudoir 173Ctenostoma (Myrmecilla) agnatum fuscum Naviaux 174Ctenostoma (Myrmecilla) albofasciatum Chaudoir 174Ctenostoma (Myrmecilla) asperulum Bates 175Ctenostoma (Myrmecilla) bicristatum Chaudoir 175Ctenostoma (Myrmecilla) brevicorne W. Horn 177Ctenostoma (Myrmecilla) breviusculum von Mannerheim 177Ctenostoma (Myrmecilla) compactum Naviaux 179Ctenostoma (Myrmecilla) coracinum Naviaux 179Ctenostoma (Myrmecilla) crucifrons W. Horn 180Ctenostoma (Myrmecilla) dentifrons W. Horn 180Ctenostoma (Myrmecilla) dokhturowi W. Horn 181Ctenostoma (Myrmecilla) flexuosum Naviaux 183Ctenostoma (Myrmecilla) gautardi Chaudoir 184Ctenostoma (Myrmecilla) globifrons W. Horn 184Ctenostoma (Myrmecilla) hirsutum W. Horn 185Ctenostoma (Myrmecilla) ichneumoneum ellipticum

Naviaux 186Ctenostoma (Myrmecilla) ichneumoneum ichneumoneum

Dejean 187Ctenostoma (Myrmecilla) infimum Naviaux 187Ctenostoma (Myrmecilla) luteum Naviaux 191Ctenostoma (Myrmecilla) macilentum Klug 191Ctenostoma (Myrmecilla) minusculum Naviaux 192Ctenostoma (Myrmecilla) modicum Naviaux 192Ctenostoma (Myrmecilla) obliquatum Chaudoir 194Ctenostoma (Myrmecilla) oblitum Chaudoir 194Ctenostoma (Myrmecilla) parvulum Naviaux 196Ctenostoma (Myrmecilla) plicaticolle W. Horn 197Ctenostoma (Myrmecilla) pygmaeum concolor Naviaux 197Ctenostoma (Myrmecilla) pygmaeum pygmaeum

(Lacordaire) 198Ctenostoma (Myrmecilla) rivalieri Naviaux 199Ctenostoma (Myrmecilla) schaumi pilosulum W. Horn 200Ctenostoma (Myrmecilla) schaumi schaumi W. Horn 201Ctenostoma (Myrmecilla) schmalzi paraguayense Brouerius

van Nidek 201Ctenostoma (Myrmecilla) schmalzi schmalzi W. Horn 201Ctenostoma (Myrmecilla) unifasciatum affine W. Horn 204Ctenostoma (Myrmecilla) unifasciatum unifasciatum

Dejean 204


Ctenostoma (Myrmecilla) vicinum Naviaux 205Ctenostoma (Myrmecilla) zerchei Naviaux 206Ctenostoma (Myrmecilla) zonata Chaudoir 206Ctenostoma (Naviauxiana) aeneum Naviaux 173Ctenostoma (Naviauxiana) arnaudi Naviaux 174Ctenostoma (Naviauxiana) cayennense Naviaux 178Ctenostoma (Naviauxiana) dormeri W. Horn 181Ctenostoma (Naviauxiana) durantoni Naviaux 181Ctenostoma (Naviauxiana) ebenina Bates 182Ctenostoma (Naviauxiana) ecuadorense Naviaux 182Ctenostoma (Naviauxiana) erwini Naviaux 182Ctenostoma (Naviauxiana) insigne Chaudoir 188Ctenostoma (Naviauxiana) longipalpe Naviaux 190Ctenostoma (Naviauxiana) maculosum Naviaux 191Ctenostoma (Naviauxiana) metallicum (Laporte de

Castelnau) 192Ctenostoma (Naviauxiana) nigrum lecourti Naviaux 193Ctenostoma (Naviauxiana) nigrum nigrum Chaudoir 193Ctenostoma (Naviauxiana) onorei colombiense Naviaux 195Ctenostoma (Naviauxiana) onorei onorei Naviaux 195Ctenostoma (Naviauxiana) ornatum ornatum Klug 195Ctenostoma (Naviauxiana) ornatum werneri Naviaux 195Ctenostoma (Naviauxiana) panamense Naviaux &

Brzoska 196Ctenostoma (Naviauxiana) pearsoni Naviaux 196Ctenostoma (Naviauxiana) sallei Chaudoir 200Ctenostoma (Naviauxiana) simile Naviaux 201Ctenostoma (Naviauxiana) spinosum Naviaux 202Ctenostoma (Naviauxiana) subtilesculptum W. Horn 202Ctenostoma (Neoprocephalus) angustoobliquatum W. Horn 174Ctenostoma (Neoprocephalus) brendelli Naviaux 176Ctenostoma (Neoprocephalus) brunneum Naviaux 178Ctenostoma (Neoprocephalus) cassolai Naviaux 178Ctenostoma (Neoprocephalus) cylindratum Naviaux 180Ctenostoma (Neoprocephalus) davidsoni Naviaux 180Ctenostoma (Neoprocephalus) deuvei Naviaux 181Ctenostoma (Neoprocephalus) germaini W. Horn 184Ctenostoma (Neoprocephalus) guatemalense Brouerius

van Nidek 185Ctenostoma (Neoprocephalus) hovorei Naviaux &

Brzoska 186Ctenostoma (Neoprocephalus) ibidion ibidion Dohrn 186Ctenostoma (Neoprocephalus) ibidion jolyi Naviaux 186Ctenostoma (Neoprocephalus) immaculatum W. Horn 187Ctenostoma (Neoprocephalus) intermedium Naviaux 188Ctenostoma (Neoprocephalus) johnsoni Naviaux 189Ctenostoma (Neoprocephalus) laeticolor Bates 189

Ctenostoma (Neoprocephalus) landolti Steinheil 190Ctenostoma (Neoprocephalus) maculicorne (Chevrolat) 191Ctenostoma (Neoprocephalus) nitidum Naviaux 194Ctenostoma (Neoprocephalus) tumidum Naviaux 203Ctenostoma (Neoprocephalus) turnbowi Naviaux 204Ctenostoma (Neoprocephalus) vairai Cassola 205Ctenostoma (Neoprocephalus) wappesi Naviaux 205Ctenostoma (Paractenosoma) corculum Bates 179Ctenostoma (Paractenosoma) parallelum Navaiux 196Ctenostoma (Paractenosoma) simpliceps W. Horn 202Ctenostoma (Salvioides) zikani W. Horn 206Cychrini Laporte de Castelnau 137Cychrus Fabricius 137Cychrus hemphillii hemphillii G. Horn 138Cychrus hemphillii rickseckeri LeConte 138Cychrus tuberculatus T.W. Harris 138

LLeistus Frölich 22Leistus (Leistus) ferrugineus (Linné) 22Leistus (Neoleistus) ferruginosus von Mannerheim 23Leistus (Neoleistus) longipennis Casey 23Leistus (Neoleistus) madmeridianus Erwin 24Loricera Latreille 69Loricera aptena Ball & Erwin 70Loricera decempunctata Eschscholtz 70Loricera foveata LeConte 70Loricera pilicornis congesta von Mannerheim 71Loricera pilicornis pilicornis (Fabricius) 71Loricera rotundicollis Chaudoir 72Loricerini LeConte 69

NNebria Latreille 24Nebria (Boreonebria) crassicornis crassicornis Van Dyke 29Nebria (Boreonebria) crassicornis intermedia Van Dyke 29Nebria (Boreonebria) frigida R.F. Sahlberg 31Nebria (Boreonebria) gouleti Kavanaugh 33Nebria (Boreonebria) gyllenhali castanipes (Kirby) 34Nebria (Boreonebria) gyllenhali lassenensis Kavanaugh 34Nebria (Boreonebria) gyllenhali lindrothi Kavanaugh 35Nebria (Boreonebria) hudsonica LeConte 35Nebria (Boreonebria) lacustris bellorum Kavanaugh 37Nebria (Boreonebria) lacustris lacustris Casey 38Nebria (Boreonebria) nivalis gaspesiana Kavanaugh 41Nebria (Boreonebria) nivalis nivalis (Paykull) 41Nebria (Boreonebria) paradisi Darlington 43

INDEX | 321

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

Nebria (Reductonebria) danmanni Kavanaugh 30Nebria (Reductonebria) darlingtoni Kavanaugh 30Nebria (Reductonebria) desolata Kavanaugh 30Nebria (Reductonebria) diversa LeConte 30Nebria (Reductonebria) eschscholtzii Ménétriés 31Nebria (Reductonebria) gregaria Fischer von Waldheim 33Nebria (Reductonebria) haida Kavanaugh 35Nebria (Reductonebria) jeffreyi Kavanaugh 36Nebria (Reductonebria) lituyae Kavanaugh 38Nebria (Reductonebria) louiseae Kavanaugh 38Nebria (Reductonebria) lyelli Van Dyke 39Nebria (Reductonebria) mannerheimii Fischer von

Waldheim 39Nebria (Reductonebria) navajo Kavanaugh 41Nebria (Reductonebria) obliqua chuskae Kavanaugh 42Nebria (Reductonebria) obliqua obliqua LeConte 42Nebria (Reductonebria) pallipes Say 43Nebria (Reductonebria) sahlbergii modoc Kavanaugh 45Nebria (Reductonebria) sahlbergii sahlbergii Fischer von

Waldheim 45Nebria (Reductonebria) sahlbergii triad Kavanaugh 46Nebria (Reductonebria) suturalis LeConte 47Nebria (Reductonebria) wallowae Kavanaugh 49Nebria (Reductonebria) zioni oasis Kavanaugh 49Nebria (Reductonebria) zioni zioni Van Dyke 50Nebriini Castelnau de Laporte 22Nippononebria Uéno 50Nippononebria (Vancouveria) altisierrae (Kavanaugh) 50Nippononebria (Vancouveria) campbelli (Kavanaugh) 51Nippononebria (Vancouveria) virescens (G. Horn) 51Notiokasiini Kavanaugh & Nègre 51Notiokasis Kavanaugh & Nègre 51Notiokasis chaudoiri Kavanaugh & Nègre 52Notiophilini Mostschulsky 53Notiophilus Dumeril 53Notiophilus aeneus (Herbst) 54Notiophilus aquaticus (Linné) 54Notiophilus biguttatus (Fabricius) 55Notiophilus borealis Harris 56Notiophilus chihuahuae Casey 56Notiophilus directus Casey 56Notiophilus intermedius Lindroth 57Notiophilus nemoralis Fall 57Notiophilus nitens LeConte 57Notiophilus novemstriatus LeConte 57Notiophilus palustris (Duftschmid) 58Notiophilus rufipes Curtis 59


Notiophilus semiopacus Eschscholtz 59Notiophilus semistriatus Say 59Notiophilus sierranus Casey 60Notiophilus simulator Fall 60Notiophilus specularis Bates 60Notiophilus sylvaticus Eschscholtz 61

OOmophron Latreille 63Omophron americanum Dejean 64Omophron baenningeri Dupuis 64Omophron dentatum LeConte 64Omophron dominicense Chaudoir 65Omophron gilae LeConte 65Omophron gratum Chaudoir 65Omophron grossum Casey 66Omophron labiatum (Fabricius) 66Omophron mexicanum Dupuis 66Omophron nitidum LeConte 66Omophron obliteratum G. Horn 67Omophron oblongiusculum Chevrolat 67Omophron ovale G. Horn 67Omophron robustum G. Horn 68Omophron solidum Casey 68Omophron sphaericum Chevrolat 68Omophron tessellatum Say 69Omophronini Bonelli 63Opisthiini Dupuis 52Opisthius Kirby 52Opisthius richardsoni Kirby 53

PPelophila Dejean 20Pelophila borealis (Paykull) 20Pelophila rudis (LeConte) 21Pelophilini Kavanaugh 20

SScaphinotus Dejean 1826 139Scaphinotus (Brennus) bullatus Van Dyke 145Scaphinotus (Brennus) cordatus (LeConte) 145Scaphinotus (Brennus) crenatus (Motschulsky) 146Scaphinotus (Brennus) cristatus (T.W. Harris) 146Scaphinotus (Brennus) interruptus (Ménétriés) 152Scaphinotus (Brennus) johnsoni Van Dyke 153Scaphinotus (Brennus) marginatus (Fischer von

Waldheim) 155

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

Dyke 149Scaphinotus (Scaphinotus) elevatus tenebricosus Roeschke 149Scaphinotus (Scaphinotus) kelloggi (Dury) 153Scaphinotus (Scaphinotus) macrogonus Bates 154Scaphinotus (Scaphinotus) mexicanus Bates 156Scaphinotus (Scaphinotus) parisianus Allen & Carlton 157Scaphinotus (Scaphinotus) petersi biedermani Roeschke 157Scaphinotus (Scaphinotus) petersi catalinae Van Dyke 157Scaphinotus (Scaphinotus) petersi corvus (Fall) 158Scaphinotus (Scaphinotus) petersi grahami Van Dyke 158Scaphinotus (Scaphinotus) petersi kathleenae Ball 158Scaphinotus (Scaphinotus) petersi petersi Roeschke 159Scaphinotus (Scaphinotus) snowi parkeri Van Dyke 162

INDEX | 323

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

Scaphinotus (Steniridia) violaceus violaceus (LeConte) 167Scaphinotus (Stenocantharis) angusticollis (Fischer von

Waldheim) 144Scaphinotus (Stenocantharis) hatchi Beer 150Scaphinotus (Stenocantharis) velutinus (Ménétriés) 165Sphaeroderus Dejean 167Sphaeroderus bicarinatus (LeConte) 168Sphaeroderus canadensis canadensis Chaudoir 168Sphaeroderus canadensis lengi Darlington 169Sphaeroderus indianae (Blatchley) 169Sphaeroderus nitidicollis brevoorti LeConte 169Sphaeroderus nitidicollis nitidicollis Guérin-Méneville 170Sphaeroderus nitidicollis schaumii Chaudoir 170Sphaeroderus stenostomus lecontei Dejean 170Sphaeroderus stenostomus stenostomus (Weber) 171Systolosoma Solier 17Systolosoma breve Solier 17Systolosoma lateritium Nègre 18Systolosomatini Erwin 17

TTrachypachini Thomson 18Trachypachus Motschulsky 18Trachypachus gibbsii LeConte 19Trachypachus holmbergi von Mannerheim 19Trachypachus slevini Van Dyke 19


Plates

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.

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 atMoolack Beach along the coast of Oregon; BOTTOM, typical landscape (Moolack Beach along the coast ofOregon) in which members of this genus dwell. Living and landscape images credit: David R. Maddison,University of Arizona, Tucson.

Plate 2. Trachypachus Motschulsky

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 foundunder stones on rocky flats beside the river. Living and landscape image credits: Roy Anderson, Belfast, NorthernIreland, U.K. and David H. Kavanaugh, California Academy of Sciences, San Francisco, respectively.

Plate 3. Pelophila Dejean

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, CountyDown, N. Ireland, 150 meters altitude; BOTTOM, Lituya Bay, Glacier Bay National Park, Alaska where adults ofL. ferruginosus Mannerheim are found under logs and on leaves and stems of Vaccinium sp. at night. Living andlandscape image credits: Roy Anderson, Belfast, Northern Ireland, U.K. and David H. Kavanaugh, CaliforniaAcademy of Sciences, San Francisco, respectively.

Plate 4. Leistus Frölich

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, NWof 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 5. Nebria Latreille

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 StanleyPark where adults of N. virescens Horn are found hiding under flakes and in cracks on bare vertical wall of trailcut. Living and landscape images credit: David H. Kavanaugh, California Academy of Sciences, San Francisco.

Plate 6. Nippononebria Uéno

Plate 7. Notiokasis Kavanaugh & Nègre

Extended focus image, adult of Notiokasis chaudoiriKavanaugh & Nègre from Montevideo, Uruguay, ABL (adultlength) 6.5 mm (in CAS). EFI credit: David H. Kavanaugh,California Academy of Sciences, San Francisco.

TOP LEFT, extended focus image, adult of Opisthius richardsoni Kirby from near Troutdal, Oregon, ABL (adultlength) 10.0 mm (ADP108781); TOP RIGHT, Opisthius richardsoni Kirby, living beetle at Kootenay River near BullRiver, 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 ofstreams in this area. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson andDavid H. Kavanaugh, California, Academy of Sciences, San Francisco, respectively.

Plate 8. Opisthius Kirby

TOP LEFT, extended focus image, adult of Notiophilus aeneus (Herbst) from near Largo, Maryland, ABL (adultlength) 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 whichmembers of the genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona,Tucson and Grace P. Servat, UNOP, respectively.

Plate 9. Notiophilus Dumeril

TOP LEFT, extended focus image of Cicindis horni Bruch, ABL (adult length) 9.5 mm (ADP108783), from SalinasGrande, 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 thisspecies dwell. Living and landscape images credit: Valeria Aschero, CRICyT, Mendoza, Argentina.

Plate 10. Cicindis Bruch

TOP LEFT, extended focus image of Omophron americanum Dejean from Plummers Island, Maryland, ABL (adultlength) 6.0 mm (ADP034029); TOP RIGHT, Omophron tessellatum Say, living beetle at Boone River near WebsterCity, IA, 305 meters altitude; BOTTOM, typical landscape (sandy beach, Lake Huron) in which members of thisgenus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and HenriGoulet, CNC, Ottawa, Ontario, respectively.

Plate 11. Omophron 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, 50meters 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 imagecredits: Roy Anderson, Belfast, Northern Ireland, U.K. and David H. Kavanaugh, California Academy ofSciences, San Francisco, respectively.

Plate 12. Loricera Latreille

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 membersof the genus dwell. Landscape image credit: Grace P. Servat, UNOP.

Plate 13. Callisthenes Fischer von Waldheim

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 membersof the genus dwell. Landscape image credit: Grace P. Servat, UNOP

Plate 14. Calosoma (Callitropa Motschulsky)

TOP, digital image, adult of Calopachys viridissimus Haury, from 10 km NW Oaxaca, Oaxaca, México, ABL (adultlength) 21.0 mm (ADP108786); BOTTOM, typical landscape (N of Oaxaca, México) in which members of thegenus dwell. Landscape image credit: Christopher J. Earle, Gymnosperm Database (http://www.conifers.org).

Plate 15. Calopachys Haury

TOP LEFT, digital image, adult of Calosoma scrutator (Fabricius) from near Snow Hill, Maryland, ABL (adultlength) 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 whichmembers of the genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona,Tucson and Grace P. Servat, UNOP, respectively.

Plate 16. Calosoma Weber

TOP LEFT, digital image, adult of Carabus serratus Say from Plummers Island, Maryland, ABL (adult length) 19.0mm (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 oncoastal tundra under debris. Living and landscape images credit: Henri Goulet, CNC, Ottawa, Ontario and DavidH. Kavanaugh, California Academy of Sciences, San Francisco, respectively.

Plate 17. Carabus Linné

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, aboveRio Cipresal, 195 meters altitude; BOTTOM, typical landscape (Mallines, Parque Nacional Puyehue at 710maltitude) 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 18. Ceroglossus Solier

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 whereadults 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, SanFrancisco, respectively.

Plate 19. Cychrus Fabricius

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 atEagle 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 20. Scaphinotus 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 atEno River State Park, NC, 135 meters altitude; BOTTOM, typical landscape (Eno River, State Park, NC, 135meters altitude) in which members of the genus dwell. Living and landscape images credit: David R. Maddison,University of Arizona, Tucson.

Plate 21. Sphaeroderus Dejean

TOP, digital image, adult of Ctenostoma erwini Naviaux, ABL (adult length) 13.5 mm (ADP000530), from BarroColorado Island, Panamá; BOTTOM, typical suspended dry leaf habitat (Tiputini Biodiversity Station, Ecuador) inwhich members of the genus dwell. Landscape image credit: Terry L. Erwin.

Plate 22. Ctenostoma Klug

Erwin 2007 Caraboidea of the Western Hemisphere (Carabidae)

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