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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 501

The Ground-Beetles of Central America (Carabidae), Part II: Notiophilini, Loricerini, and Carabini

Terry L. Erwin

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1991

A B S T R A C T

Erwin, Terry L. The Ground-Beetles of Central America (Carabidae), Part II: Notiophilini, Loricerini, and Carabini. Smithsonian Contributions to Zoology, number 501, 30 pages, 22 figures, 5 tables, 1991.—This paper is the second in a planned series designed to cover the ground-beetles of Central America both taxonomically and biogeographically. The area, for purposes of this series, extends from the northern border of Guatemala south to the border between the Republic of Panama and Colombia, including the near shore islands of both oceans. Species that are known to occur now in either Mexico or Colombia in the vicinity of the borders shared with Central American countries, and that may someday be found in Central America, are discussed as well. Included here is part of the subfamily Carabinae, tribes Notiophilini, Loricerini, and Carabini. Other tribes of Carabinae will be covered in another fascicle. Also included herein is a key to the presently described genera of the New World tropics, a classification scheme of the world fauna, and introductory material pertinent to both the series and the present contribution. The following genera are discussed: Notiophilus, Loricera, and Calosoma. All species, none of which are new, are redescribed and some structural parts of each are illustrated. Notes are provided on natural history and distribution for each species; dot maps illustrate the known Central American range of each taxon; and line or shaded drawings depict diagnostic characters.

S U M A R I O

Este fascículo es el segundo de una serie planeada con el fin de cubrir los carábidos de América Central, tanto taxonómica como bio-geograficamente. El área en relación a esta serie se extiende desde la frontera none de Guatemala hacia el sur en el area limitrofe entre Panamá y Colombia, incluyendo las islas cercanas a las costas de ambos oceanos. También son consideradas las especies que han sido encontradas en Méjico o Colombia, muy cerca a las fronteras compartidas con los países de Aménca Central y que alguna vez podrían ser halladas en esta zona. Se incluye en la Parte I la sub-familia Carabinae, las tribus Notiophilini, Loricerini, y Carabini. Otras tribus de la misma sub-familia serán tratadas en la Parte II. iambién se incluye en la Parte II una clave para la genera del Nuevo Mundo hasta ahora descrita, un esquema de clasificación de la fauna mundial y material y métodos pertinentes a ambas series. En la Parte I, la siguiente genera es tratada: Notiophilus, Loricera, y Calosoma. Todas las especies aunque descritas anteriormente, son redescritas y algunas partes estructurales ilustradas. Se dan a conocer notas sobre historia natural y distribución para cada taxon; mapas de localizamiento ilustran el alcance de distribución de cada especie en América Central y diferentes dibujos representan caracteristicas especificas a cada una.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Monlaslrea cavernosa (Linnaeus).

Library of Congress Calaloging-in-Publication Data
Erwin, Tiny L., 1940–
The ground-beetles of Central America (Carabidae) / Terry L. Erwin.
p. cm. - (Smithsonian contributions to zoology ; no. <501>)
Abstract in English and Spanish.
Includes bibliographical references.
Contents: - pt. 2. Notiophilini, Loricerini, and Caribini.
Supt of Docs. no.: SI 1.27:501

1. Carabidae-Central America-Classification. 2. Carabidae-Central America-Geographical distribution. I. Title. II. Series.

QL1.S54 no. 501, etc.

[QL596.C2]

591 s-dc20

90-10363

[595.76′2]

CIP

Contents

Introduction

Acknowledgments

Methods

Materials

Key to Tribes and Some Genera of Neotropical Carabidae

Classification of Caraboidea

Tribe NOTIOPHILINI

Checklist of Notiophilini of Central America

Genus Notiophilus Duméril

Notiophilus specularis Bates

Tribe LORICERINI

Checklist of Loricerini of Central America

Genus Loricera Latreille

Loricera rotundicollis Chaudoir

Tribe CARABINI

Checklist of Carabini of Central America

Genus Calosoma Weber

Key to the Calosoma of Central America

Calosoma fulgens Chaudoir

Calosoma alternans (Fabricius)

Calosoma sayi Dejean

Calosoma abbreviatum Chaudoir

Calosoma scrutator (Fabricius)

Calosoma aurocinctum Chaudoir

Calosoma angulatum Chevrolat

Calosoma marginalis Casey

Calosoma glabratum Dejean

Calosoma ampliator Bates

Literature Cited

REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION

Dedication

To those who saw the forests before and marveled at their beauty, to those who knew not mass extinction, and to those who still appreciate a mountain, a brook, a beetle; especially to my professor and mentor George E. Ball, upon his retirement from the formality of a classroom; may he never give up teaching.

The Ground-Beetles of Central America (Carabidae), Part II: Notiophilini, Loricerini, and Carabini

Terry L. Erwin

____________

Terry L. Erwin, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560.

Introduction

One day long ago, George E. Ball asked me to accompany him on a field trip to Mexico. I was a graduate student and had never been to the tropics; I was obviously excited! We were collecting carabids by hand each day and by light on most nights. While the ultraviolet collecting lamp was running, we prepared the previous day’s catch in the back of George’s “El Escarabajo-I” (a large well-stocked camper truck). George wrote up the notes and I struggled to sort all the collected specimens to the morpho-species level for him. Two months of personal tutoring from George during that trip gave me tropical carabid fever and the initial background necessary to devote my career to the fascinating Neotropics and the wealth of hidden evolutionary treasures dwelling there. Since that first introduction, 16 years of organizing specimens (over 100,000) and their accompanying data, studying types, and accumulating literature has taken place. As a result, this fascicle, actually the second of a planned series (see Erwin, 1982a), which will cover faunistically all Central American carabid species, begins to provide a systematic foundation for future ground-beetle studies not only of Central America, but of the West Indies and South America as well. This series then, I dedicate to George—his enthusiasm, his inspiration, and his leadership.

Carl Lindroth’s monumental, 6-part “Ground-Beetles of Canada and Alaska” (1961–1969) appeared beginning with Part 2. Darlington’s equally important “Carabid Beetles of New Guinea” (1952–1971) began with Part 2 also, although for different reasons. In any large-scale serialized monographic treatment of a major fauna, one never knows initially what to include in the preliminary or introductory part Lindroth published the taxonomy first (Parts 2 through 6) then provided Part 1 (1969), which described methods, procedures, and a historical account of work done on the group for the area of coverage. Darlington, on the other hand, began with his favorite taxon, the Agonini, and provided a quite complete introductory foreword for it and all the subsequent parts, although he also wrote smaller, separate introductions to each of them too. I have chosen here to provide a terse introduction, then proceed with the taxonomy. A complete separate introduction and analysis will be published last. This last part will include methods, procedures, and other standard items (e.g., see Erwin, 1970, 1973, 1974, 1975, 1982a); however most of these have been covered in one way or another in my previous papers.

Because the most important first step in studying any fauna or flora is identification of the taxa, I herein provide a provisional key to all tribes and some genera known presently in the fauna. A more extensive key will accompany the last part. Arrangement of taxa follows that of Erwin (1985). New taxa and additional data pertaining to species covered herein, but discovered after this volume was prepared, will be included in a later supplement. Taxa that I predict may be found in the fauna with additional collecting are keyed and described, but these are not numbered as are the known Central American species.

Lindroth (1969) provided useful data for those becoming acquainted with ground beetles, and these data are especially applicable here. For a general synthesis of knowledge about ground-beetles, the reader is referred to Erwin et al. (1979); for more specifics concerning Neotropical Carabidae see Reichardt (1977) and Erwin and Sims (1984).

The main purpose of this coverage of Central American ground-beetles is to provide the user with an efficient yet simple tool for identification of taxa. It will also lead the user to an accumulated digest of all that is presently known about the taxa, including ecologic and geographic ranges, phylogenetic relationships, and natural history.

Acknowledgments

There is little doubt that I could not have begun this series of papers without many discussions with my mentor and friend George E. Ball, with whom such discussions still continue, and to the late Carl H. Lindroth and Philip Darlington Jr., my mentors earlier in my career, who set such a fine examples with their faunal treatments discussed below. I also acknowledge with great appreciation the interchanges, both verbal and written, with David H. Kavanaugh, Gerald R. Noonan, Nigel E. Stork, and Donald R. Whitehead. I thank, for the present paper, George Venable, Gloria Gordon Zimmer, and Sophie Allington for the illustrations, Linda Sims for general assistance in preparation of the manuscript and management of the computerized data base, Gloria N. House for managing the specimens, Leslie Schimmel and Jeannine Weaver for measuring them, Dora Rios for translation of the abstract to Spanish, and the Fluid Research and Environmental Sciences Programs of the Smithsonian Institution and the National Museum of Natural History for providing funding for extensive field work and museum visits over the years. James Liebherr, Lee Herman, Ronald McGinley, and Donald Whitehead provided excellent reviews of the manuscript and contributed significantly to its final form and for that I am grateful, but I take full responsibility for any remaining problems should there be any.

I also very much appreciate the efforts made by several museum curators who loaned material or allowed me to study type specimens in their charge: L. Herman, American Museum of Natural History, New York (AMNH); British Museum (Natural History), London, England (BMNH); David H. Kavanaugh, California Academy of Sciences, San Francisco, California (CAS); Ronald D. Cave, Department of Entomology, Auburn University, Auburn, Alabama (CAVERD); Robert Woodruff, Florida Department of Agriculture, Gainesville, Florida (FDAG); L. Watrous (retired), Field Museum of Natural History, Chicago, Illinois (FMNH); GR. Noonan, Milwaukee Public Museum, Milwaukee, Wisconsin (GRNO); J. Nègre (deceased), 78 Versailles, 9, Bb, De Lesseps, France (JNEG); A. Newton (now at FMNH), Museum of Comparative Zoology, Cambridge, Massachusetts (MCZ); H. Perrin, J. Menier, Muséum National d'Histoire Naturelle, Paris, France (MNHP); C.A. Triplehom, Department of Entomology, Ohio State University, Columbus, Ohio (OSU); Henry P. Stockwell, Gorgas Memorial Hospital, Canal Zone, Republic of Panama (STOCHP); George E. Ball, Strickland Museum, University of Alberta, Edmonton, Canada (UASM); R.T. Allen, University of Arkansas, Fayetteville, Arkansas (UAIQ; R. Fisher, University of Michigan, Ann Arbor, Michigan (UMA A); and Department of Entomology, Smithsonian Institution, Washington, D.C. (USNM).

Methods

General procedural methods for the series are those that I used before (Erwin, 1970, 1973, 1974, 1975; Erwin and Kavanaugh, 1981). Measures for various body dimensions are coded as follows and are presented in the species descriptions as single specimen measures (if that is all I saw) or as ranges based on the smallest and largest of all specimens studied. All specimens were measured with an electronic measuring device (Erwin, 1978) and measures are presented in millimeters. ABL = apparent body length, that length used by most previous authors as total length as measured by holding up a ruler alongside the specimen (see Erwin and Kavanaugh, 1981). TW = total width across the widest portion of the elytra, actually measured as the left elytron (WE) and doubled to obtain value. All label, specimen measurement, and field data about specimens are computerized (LJ.M. Erwin, 1976), and these records formed the base for study and analysis. Data concerning all Central American material examined are presented in tables and maps (ranges are for Central America only); tables summarize data deduced from specimen labels. In the tables, most localities are presented as latitude/longitude junctures; but for type specimens, place names are cited in the text. Elevations are given in meters even though specimen labels may have them in English units; occasionally, specimen labels give imprecise altitudinal data or none at all, thus in these cases tables include the approximation symbol (~) to indicate approximate altitude derived from topographic maps. When known, a descriptor of the general habitat is given in terms of the Holdridge classification of the Central American environment. Under “Natural History” notes, below, the Holdridge life zones are capitalized. Microsculpture micrographs will be made only for species where sufficient study material allowed coating of body parts and where such character states are necessary for identification.

At present, a problem exists in accurately describing colors in ground-beetles, especially those hues ranging from white-yellow through brown to black. Blues, reds, and greens can, more or less, be described using the English words. Until such time as an international color standard chart is adopted, I am using the following Latin-based terms: BROWN—brunneous, dusky dark brown with blackish overtones; castaneous, rich chestnut; testaceous, brownish yellow. YELLOW—flavous, golden yellow; fulvous, reddish yellow. RED—rufous, reddish brown; ferrugineous, rust red. BLACK—fuscous, dusky black with brown overtones; piceous, black with reddish overtones; and ebineous, pure black.

As in all large-scale faunal coverages, new material constantly trickles in to the scientist doing the study. Such material received by me after the completion of each fascicle will be covered in a series of supplements. The main series will be published in the present monographic serial; thus future binding of the entire series will be facilitated by a constant style and format

For ease of finding taxa throughout the series, a numbering system is used as in Erwin (1982b). Tribes are given a number (00) in the list of classified tribes below. Within each tribe, each genus is given a number (.00) and within genera, each species (.000). Reference between keys and descriptions is thus facilitated.

Materials

This paper is based on examination of 78 specimens from Central America. I also studied numerous specimens from adjacent areas, of Mexico and Colombia, as well as numerous specimens from other parts of Latin America. These specimens are not formally listed, but are covered under the heading “Geographical Distribution” under each species description. Under “Acknowledgments” are listed the depositories for these materials. Acronyms for personal or institutional collections based on Erwin (1970) are given following each entry.

Key to Tribes and Some Genera of Neotropical Carabidae

(Modified from G.E. Ball in Reichardt, 1977)

Notes on Use of Keys: The following key-steps lead either to tribes with a single listed genus (for which discussions can be found in the following pages) or to tribes with several genera. In the latter case, these genera are keyed elsewhere (for the location, see Contents and previous or subsequent parts of the series). Because it is possible that groups occurring in Mexico and South America may yet be discovered in Central America, I have presented a key here to all tropical groups now known from the Western Hemisphere. Illustrations for general carabid characteristics can be found in Ball (1960) and Lindroth (1969) and will be elaborated in the faunal fascicle of this series.

Classification of Caraboidea

(Italicized taxa occur in Central America)

SUPERFAMILY CARABOIDEA

I. Family Trachypachidae

01. Tribe Trachypachini

02. Tribe Systolosomini

II. Family Carabidae

Division Nebriiformes

A. Subfamily Carabinae

a. Supertribe Nebriitae

01. Tribe Nebriini

02. Tribe Notiokasiini

03. Tribe Opisthiini

04. Tribe Cicindisini

05. Tribe Notiophilini

b. Supertribe Loriceritae

06. Tribe Loricerini

c. Supertribe Carabitae

07. Tribe Carabini

08. Tribe Ceroglossini

09. Tribe Pamborini

10. Tribe Cychrini

d. Supertribe Cicindelitae

11. Tribe Collyrini

12. Tribe Megacephalini

13. Tribe Ctenostomatini

14. Tribe Mantichorini

15. Tribe Cicindelini

e. Supertribe Omophronitae

16. Tribe Omophronini

Division Loxomeriformes

B. Subfamily Scaritinae

f. Supertribe Migadopitae

17. Tribe Amarotypini

18. Tribe Migadopini

g. Supertribe Elaphritae

19. Tribe Elaphrini

h. Supertribe Promecognathitae

20. Tribe Promecognathini

i. Supertribe Siagonitae

21. Tribe Enceladini

22. Tribe Siagonini

j. Supertribe Hiletitae

23. Tribe Hiletini

k. Supertribe Pseudomorphitae

24. Tribe Pseudomorphini

l. Supertribe Scarititae

25. Tribe Cnemacanthini

26. Tribe Scaritini

27. Tribe Clivinini

C. Subfamily Paussinae

m. Supertribe Metriitae

28. Tribe Metriini

n. Supertribe Paussitae

29. Tribe Nototylini

30. Tribe Mystropomini

31. Tribe Ozaenini

32. Tribe Protopaussini

33. Tribe Paussini

o. Supertribe Brachinitae

34. Tribe Crepidogastrini

35. Tribe Brachinini

Division Melaeniformes

D. Subfamily Broscinae

p. Supertribe Melaenitae

36. Tribe Melaenini

37. Tribe Cymbionotini

q. Supertribe Broscitae

38. Tribe Broscini

r. Supertribe Apotomitae

39. Tribe Apotomini

Division Psydriformes

E. Subfamily Psydrinae

s. Supertribe Psydritae

40. Tribe Gehringiini

41. Tribe Psydrini

42. Tribe Melisoderini

43. Tribe Tropidopterini

44. Tribe Meonidini

55. Tribe Patrobini

46. Tribe Amblytelini

t. Supertribe Rhysoditae

47. Tribe Rhysodini

u. Supertribe Trechitae

48. Tribe Trechini

49. Tribe Zolini

55. Tribe Pogonini

51. Tribe Bembidiini

F. Subfamily Harpalinae

v. Supertribe Pterostichitae

52. Tribe Morionini

53. Tribe Pterostchini

54. Tribe Zabrini

w. Supertribe Panagaeitae

55. Tribe Bascanini

56. Tribe Panagaeini

57. Tribe Agonicini

58. Tribe Disphaericini

59. Tribe Peleciini

x. Supertribe Callistitae

60. Tribe Cuneipectini

61. Tribe Callistini

62. Tribe Chaetogenyini

63. Tribe Oodini

64. Tribe Licinini

y. Supertribe Harpalitae

65. Tribe Harpalini

z. Supertribe Dryptitae

66. Tribe Dryptini

67. Tribe Zuphiini

68. Tribe Galeritini

a′. Supertribe Anthiitae

69. Tribe Helluonini

70. Tribe Anthiini

71. Tribe Helluodini

b′. Supertribe Orthogoniitae

72. Tribe Idiomorphini

73. Tribe Amorphomerini

74. Tribe Orthogoniini

75. Tribe Catapiesini

c′. Suptertribe Ctenodactylitae

76. Tribe Hexagoniini

77. Tribe Ctenodactylini

d′. Supertribe Platynitae

78. Tribe Platynini

e′. Supertribe Lebiitae

79. Tribe Perigonini

80. Tribe Lachnophorini

81. Tribe Graphipterini

82. Tribe Cyclosomini

83. Tribe Masoreini

84. Tribe Odacanthini

85. Tribe Lebiini

05 Tribe NOTIOPHILINI

The notiophilines are highly distinctive terrestrial beetles. They have a large head relative to the narrow prothorax, huge hemispherical eyes, and numerous frontal carinae; many species possess shiny elytral intervals as well. These small beetles are predominantly Holarctic in distribution, with the exception of the single Central American species, which is widespread in Mexico and occurs in Guatemala and El Salvador only at higher elevations. The group is ancient and contains presendy but one genus, Notiophilus. As in the loricerines, amarines, and trechines, this group is but a small element in the Central American fauna; however, the species is important in zoogeographic perspective because it represents a southward extension of a widespread temperate group.

The notiophilines, by most analyses, are primitive carabids; however, because of their unique body form and peculiar character states, it has not been possible to determine sister group relationships. The male parameres are nebrioid, but the median lobe is as peculiar as the external habitus. A single clue to the relationship of these beetles might lie in an observed, but so far unnoted character state of the middle tarsal articles. Adults of Omopkron, Trachypachus, Notiophilus, and Hiletini have the basitarsomere of the middle legs slightly dilated with squamate setae beneath. This feature is apotypic in the family and occurs nowhere else among the more primitive lineages. The question is whether it is synapotypic or convergent.

For the most recent treatment of this tribe, see Lindroth (1961).

Checklist of Notiophilini of Central America

05.01 Notiophilus Duméril (1806:194)

05.01.001 N. specularis Bates (1881:19)

05.01 Genus Notiophilus Duméril

Notiophilus Duméril, 1806:194. [Type species, Carabus aquaticus Linné, 1758:408; subsequent designation by Westwood, 1840, 2:6.]

DIAGNOSTIC COMBINATION.—Small to medium-small beetles, somewhat depressed and rectangulate in form; forebody and head broad; appendages slender and of medium length in proportion to body, eyes large. Color black with brassy luster, often appendages or part of elytron paler. Frons with series of longitudinal carinae. Middle coxal cavities conjunct (entirely enclosed by sterna). Size small, length less than 7.0 mm.

NATURAL HISTORY.—Notiophilus usually live in forested habitats among conifer needles, mosses, or in broadleaf tree litter. Some species have adapted to gardens around human residences. Adults are carnivorous, feeding on gnats and spiders, which they capture during their diurnal activities; larvae are also carnivorous, but their diet needs study. The huge eyes of adults, like tiger beetles and certain log-running carabids, are adapted for visual hunting. Most species are “heliophilous and rather xerophilous” (Lindroth, 1961:91). Although wing dimorphism is common in the more northern species, N. specularis are apparently always fully winged. Larval stages are known for two northern species but, according to Lindroth (1961:91), “most species probably hibernate as adults.”

In the New World there are 16 species belonging to this genus, only one of which reaches Central America in the northern most portion. Its characteristic habitus (Figure 1) makes it easily identified, even in the field. The males have three narrowly dilated basal anterior tarsal articles with squamate setae beneath, and one basal middle tarsal article also with squamate setae. In addition, male palpi are generally more dilated than in the females, thus sex determination is easily done with a hand lens in the field.

FIGURE 1.—Habitus of Notiophilus specularis Bates, dorsal aspect, female from 12.8 mi (20.6 km) E Sta. Catarina Juquila, Oaxaca, Mexico.

TAXONOMY.—Casey (1913:47) described Notiophilus chihuahuae from the Sierra Madre Mountains, Chihuahua, Mexico (lectotype, a male, No. 46839, herein selected, in USNM). This species is apparently valid and keys to N. biguttatus in Lindroth (1961) except that individuals are concolorous. It is unrelated to the more southern N. specularis, which appears to be sister species of N. obscurus Fall of California’s Sierra Nevada and San Bernardino mountains.

NOTES.—Lindroth (pers. comm.) stated that he often entertained the notion to work on this interesting genus, but it would require great amounts of material because of intraspecific variation over the range of wide-ranging species. He never got around to the task, but it should be done, for the biogeography of this ancient lineage might prove instructive, as well as interesting, in shedding light on early Tertiary events.

05.01.001 Notiophilus specularis Bates

FIGURES 1-3

Notiophilus specularis Bates, 1881:19. [Lcctotype female, MEXICO, Vera Cruz, Las Vigas, (Höge)(BMNH), herein selected.]

DIAGNOSTIC COMBINATION.—Frons with series of longitudinal costae, eyes large and hemispherical. Color and luster: black with metallic luster, appendages rufopiceous except testaceous venter of antennal articles 1 and 2. Form: size small, head very broad, eyes large, body flat. Structure: dorsal microsculpture isodiametric and granulate, absent from disc of pronotum and elytra, these very shiny. Male median lobe (Figure 2). Female styli small, trianguloid, with ventral seta. ABL = 5.3 to 6.5 mm; TW = 1.3 to 2.4 mm.

GEOGRAPHICAL DISTRIBUTION (Figure 3).—The range of this species extends from the highlands of middle Mexico south to El Salvador.

FIGURE 2.—Medium lobe of male genitalia, left lateral aspect, of Notiophilus specularis Bates, from 21.8 mi (35.1 km) N Juchatengo, 7100' (2164 m), Oaxaca, Mexico.

NATURAL HISTORY.—Notiophilus specularis is generally found, in Mexico, in pine-oak forests and cloud forests above 2000 m, and has been found in El Salvador at 2300 m.

NOTES.—The Central American species is unique among all other carabids in the fauna in having the disc of the elytra shiny and mirror-like contrasted against dull marginal areas. As in the following tribe, these beetles are not likely to be confused with any others in Central America.

MATERIAL EXAMINED.—Type (see above), 9 paralectotypes (BMNH), and 25 specimens from Mexico (UASM), 2 from El Salvador (USNM); one of the paralectotypes is from San Geronimo, Guatemala (BMNH) (Table 1).

06 Tribe LORICERINI

The loricerines are a lineage of medium-sized and peculiar ground beetles whose autapotypic features include numerous thickened setae on the antennae and mouthparts and 12 striate elytral interneurs. Most species are north temperate in distribution, except the single species discussed below, whose range extends into the Neotropical region at middle to high altitudes. A few other species occur in temperate habitats in the northern portions of the Oriental Region. The group is ancient and contains presently but one genus, Loricera, which contains two subgenera. As in the notiophilines, amarines, and trechines, this group is but a small element in the Central American fauna; however, the species is important in zoogeographic perspective because it represents a southward extension of a widespread temperate group.

The loricerines, by most analyses, are primitive carabids; however, because of their unique and peculiar character states, Ball and Erwin (1969) did not discover the sister group of Loricerini nor did they place the loricerines in any new place in the carabid classification of the time. I can do little better today. Although it is generally agreed that the group is ancient and related to the caraboid/nebrioid lineages (the male parameres are nebrioid, but the median lobe is peculiar in that it resembles the form found in more derived carabids), it must remain a group unto itself, and perhaps that is where it belongs, as a sister to the caraboid lineage.

For recent treatments of this tribe, see Lindroth (1961) and Ball and Erwin (1969).

Checklist of Loricerini of Central America

06.01 Loricera Latreille (1802:88)

06.01.001 L. rotundicollis Chaudoir (1863:115)

06.01 Genus Loricera Latreille

loricera Latreille, 1802:88. [Type species: Carabus pilicornis Fabricius, 1775:243. Monotypy.]

See Ball and Erwin, 1969, for species groups and another Old World subgenus.

DIAGNOSTIC COMBINATION.—Antennomeres 2–6 with markedly large setae; antennomeres 2–4 irregular in shape. Head with two large foveae and deep transverse sulcus behind eyes. Elytron with 12 regular striate interneurs.

FIGURE 3.—Geographical distribution map of Notiophilus specularis Bates in Central America.

TABLE 1.—Central American locality data deduced from specimen labels; 05.01.001 Notiophilus specularis Bates; see Figure 3.

FIGURE 4.—Habitus of Loricera rotundicollis Chaudoir, dorsal aspect, male from 27.0 km NW San Marcos, 2800–2900 m, San Marcos, Guatemala.

NATURAL HISTORY.—This genus is composed of 10 species, only one of which enters Central America in the north. Its characteristic habitus (Figure 4) makes it easily identified, even in the field. The males have three quite broadly dilated basal anterior tarsal articles with squamate setae beneath, thus can be easily recognized as males in the field.

The peculiar setal patterns of the adult head, mouthparts, and antennae indicate a specialized predaceous mode of living, however their prey is unknown. Adults are nocturnal and can be found at night walking over the surface of the soil or litter. During the day, individuals hide under the litter or forest floor debris. Larvae have been found under large stones.

NOTES.—Although Ball and Erwin (1969) recently reviewed this group, they did not fully revise it, determine the sister group, nor plot actual distributions of the species. These tasks remain to done, especially for the Old World.

06.01.001 Loricera rotundicollis Chaudoir

FIGURES 4-6

Loricera rotundicollis Chaudoir, 1863:115. [Lectotype female, MEXICO, Oaxaca, Capulapam, (Sallé) (MNHP), herein selected.]

DIAGNOSTIC COMBINATION.—Antennomeres 2–6 with markedly large setae; antennomeres 2–4 irregular in shape; elytron with 12 regular striate interneurs. Color and luster: mostly black, shining iridescent except head, with rufous or rufopiceous labrum, antennal articles 2, 4-11, mouthparts except palpi, coxae, trochanters, and tibiae; tarsal articles rufoflavous; palpi flavous, antennal articles 1 and 3 piceous. Form: size medium, head round with constricted neck, eyes small but prominent, body flat. Structure: dorsal microsculp ture of small, well-impressed transverse meshes. Male median lobe (Figure 5). Female styli small and spatulate with single ventral setae; coxite densely setiferous. ABL = 7.5 to 8.2 mm; TW = 2.9 to 3.4.

FIGURE 5.—Medium lobe of male genitalia, left lateral aspect, of Loricera rotundicollis Chaudoir, left lateral aspect, male from 27.0 km NW San Marcos, 2800-2900 m, San Marcos, Guatemala.

GEOGRAPHICAL DISTRIBUTION (Figure 6).—The one species is widespread in Mexico from the southern portion of the Sierra Madre Oriental south through Chiapas; in Central America, it currently is known only from the northwest portion of Guatemala, at middle to high elevation.

NATURAL HISTORY.—In Mexico, Loricera rotundicollis is generally found in oak and pine forests above 6000 feet in montane moist forest, sometimes as high as 11,000 feet. I expect that this will also be true for the northern parts of Guatemala, although this species is now known from only a few scattered localities there. The larvae are probably specialized predators and can be found under large stones where they also pupate (see Ball and Erwin, 1969, for descriptions of immature stages). All specimens seen by me from Guatemala are brachypterous, with the flight wing developed only to the stigma. Ball and Erwin (1969) point out that of 131 specimens seen by them, 119 were brachypterous and that fully winged individuals were found only in large population samples. I have no large samples from Guatemala at present, but expect winged individuals to be found there.

MATERIAL EXAMINED.—Type (see above) and 6 specimens (Table 2).

07 Tribe CARABINI

The carabines are a numerous and diverse group of small to large beetles found in most regions of the world. The male genitalia of the Carabini, sensu stricto, Cychrini, Pamborini, Ceroglossini, the so-called “Cicindelinae,” and several other minor groups are virtually identical in their ground plan structure. In addition, these groups possess the primitive antennal comb and thoracic structures, as well as vertically oriented female gonocoxites (apotypically, carabid gonocoxites are horizontally oriented) of early carabid evolution. These groups form the Carabinae, of which only two subgroups are found in Central America—Carabini (Calosomd) and Cicindelitae. The former is covered below, while I shall treat the latter in another fascicle of the series. Many authors (Breuning, 1927–1931; Jeannel, 1940; Gidaspow, 1959, 1963; Lindroth, 1961) have discussed parts of this large group, but as yet no one has dealt with generic relationships in a modern way, for the world. Although this task would be horrendous because of excessive taxonomic splitting in Carabus, it is important that it be done because of the subfamily’s interesting, and perhaps instructive, distribution.

The carabine complex alone should elucidate patterns of continental drift with regard to early carabid dispersal patterns. Jeannel (1940) discussed Calosoma distribution in relation to shifting continents, but he presented no clear evidence of group relationships, although the essence of his conclusions is probably very good. This plateau of carabid evolution was probably the last or next to last major pre-rift radiation in the family (Erwin, 1979, 1985; Kavanaugh and Negre, 1983). Inclusive taxa derived from this plateau dispersed perhaps across relatively narrow water gaps, but nevertheless were subject to an oceanic filter effect. Still later groups arose, dispersed in the fashion described by Darlington (1957) and Erwin (1985), and gave us the complex caraboid distribution pattern seen today.

FIGURE 6.—Geographical distribution map of Loricera rotundicollis Chaudoir in Central America.

TABLE 2.—Central American locality data deduced from specimen labels; 06.01.001 Loricera rotundicollis Chaudoir; see Figure 6.

Checklist of Carabini of Central America

07.01     Calosoma Weber (1801:20)

Castrida Motschulsky (1865:300)

—(C. fulgens Chaudoir, 1869:370)

07.01.001 C. alternans Fabricius (1792:146)

07.01.002 C. sayi Dejean (1826:198)

—(C. abbreviation Chaudoir, 1869:371)

Calodrepa Motschulsky (1865:310)

07.01.003 C. scrutator Fabricius (1775:239)

07.01.004 C. aurocinctum Chaudoir (1850:420)

Carabosoma Géhin (1885:32)

07.01.005 C. angulatum Chevrolat (1834, fasc. 2:44)

Camegonia Lapouge (1924:38)

07.01.006 C. marginalis Casey (1897:340)

Camedula Motschulsky (1865:303)

07.01.007 C. glabratum Dejean (1831:565)

Chrysostigma Kirby, 1837:19

07.01.008 C. ampliator Bates (1891:223)

07.01 Genus Calosoma Weber

Calosoma Weber, 1801:20. [Type species: Carabus sycophanla Linné, 1758:414; subsequent designation by Latreille, 1810:426.]

Chrysostigma Kirby, 1837:19. [Type species: Carabus calidum Fabricius, 1775:237.]

Castrida Motschulsky, 1865:300. [Type species: Calosoma sayi Dejean, 1826:198.]

Camedula Motschulsky, 1865:303. [Type species: Calosoma glabratum Dejean. 1831:565.]

Calodrepa Motschulsky, 1865:310. [Type species: Calosoma scrutator Fabricius, 1775:239.]

Carabosoma Géhin, 1885:32. [Type species: Calosoma angulatum Chevrolat, 1834, fasc. 2:44.]

Camegonia Lapouge, 1924:38. [Type species: Calosoma prominens LeConte, 1853:400.]

See Gidaspow (1959, 1963) and Lindroth (1961) for complete synonymies and modem revisions of the New World species. Old World nomenclature is not covered here.

DIAGNOSTIC COMBINATION.—Very large, robust beetles with broadly inflated elytra and narrow forebody (Figure 7). Color somber black or piceous, or metallic often with contrasting marginal stripes and/or elytral foveae; appendages piceous or black, or with faint metallic reflections if not solid metallic. Mandibles unisetose. Antennal comb terminal, transverse, between slightly offset terminal spurs. Anterior coxae open behind; middle coxae disjunct; hind coxae contiguous. Elytra with 15 or 16 intemeurs, often obliterated by macrosculpture; intervals 4, 8, and 12 with several, often metallic, foveae. All Central American species fully winged. Anterior tarsal articles of male 1 + 2, or 1 − 4 dilated and pubescent beneath. Male parameres asetose, nearly symmetric. Female with spatulate, ventrally setiferous styli; coxites densely setiferous; ovipositor oriented vertically. ABL = 18.5 to 35.0 mm in Central American species.

NATURAL HISTORY.—The species of this genus are diverse in their habitat preference. Many species occur in open woodland, hardwood forests, prairie, acacia scrub, and riverine forests. They are highly vagile and move around to areas of caterpillar outbreaks where both adults and larvae feed ravenous on the caterpillars. Many Calosoma are semiarboreal and will climb the trees for their meal; the larvae are not generally good climbers, however. Adults hibernate and often live more than one year. Sex of adults can be determined by reference to the anterior tarsal articles, some of which will be dilated and pubescent beneath in males.

Key to the Calosoma of Central America

(adapted from Gidaspow, 1963)

Calosoma fulgens Chaudoir

FIGURES 8. 10

Calosoma fulgens Chaudoir, 1869:370. [Holotype male, PARAGUAY (Bonpland) (MNHP), as originally given by Chaudoir.]

DIAGNOSTIC COMBINATION.—Color a cupreous sheen with non-contrasting elytral foveae in intervals 3, 7, and 11; middle tibia markedly arcuate; hind angle of pronotum setigerous. Color and luster: dark, metallic green, disc of elytra bright cupreous, mouthparts, appendages, and most of the venter dark piceous, metapistemum with metallic reflections. Form: medium-sized and narrow, females with apparently inflated elytra; eyes large and prominent. Structure: elytral intervals moderately convex; head finely punctulate on disc of frons, more coarsely punctate and rugose laterally. Male with 3 dilated basal protarsal articles each with setiferous pad beneath, metatrochanter ventrally hooked apically, and with median lobe sharply bent laterally near apex. Female without tarsal pads, trochanter hooks, but with multiple setae on sternum VI. Microsculpture mostly effaced from dorsal surface, finely isodiametric where present ABL = 24.0 to 28.0 mm; TW = 10.0 to 11.0 mm (from Gidaspow, 1963).

GEOGRAPHICAL DISTRIBUTION.—Gidaspow (1963) stated that these beetles are distributed from Paraguay to Colombia.

NATURAL HISTORY.—According to Gidaspow (1963), this species occurs in grasslands, prairies, and savannas amongst herbaceous vegetation. This kind of habitat results from agricultural practices of mankind, and it is likely that species occurring in northern South America will invade Panama as the forests are cut away to make pastures and croplands. Calosoma fulgens is known from Cali, Colombia, and should be looked for in eastern Panama and on the Azuero Peninsula.

MATERIAL EXAMINED.—Type (see above), 3 specimens from Ecuador (USNM).

07.01.001 Calosoma alternans (Fabricius)

FIGURES 11, 20

Carabus alternans Fabricius, 1792:146. [Holotype in ZMC according to Gidaspow (1963).]

Carabus armatum Reiche, 1842:377. [Lectotype, male, in MNHP, herein selected.]

Callistriga caxale Motschulsky, 1865:307. [New name for Calosoma armatum Reiche. 1842:377 (not armata Laporte, see below).]

DIAGNOSTIC COMBINATION.—These beetles are at once recognized by their dull, blackish elytra with alternating rows of wide and narrow intervals, the 3, 7, and 11 each with large contrasting foveae. In addition, the middle tibia is markedly arcuate and the basally lobed pronotum has latero-basal setae, one near each hind angle. Color and luster: blackish with faint cupreous reflections, somewhat greenish on head and pronotum, mouthparts, appendages and venter dark piceous. Form: medium-sized and narrow, females with more apparendy inflated elytra; eyes large and prominent. Structure: elytral intervals convex, 3, 7, and 11 catenate with large deep foveae; all intervals with transversely arranged scales; head with moderately coarse punctulae. Microsculpture finely isodiametric, effaced from surface of pronotum. Male with 3 dilated basal protarsal articles each with setiferous pad beneath, middle tibia with small brush of setae apicomedially, metatrochanter ventrally hooked apically, and median lobe moderately bent laterally near narrowed apex. Female without tarsal pads, trochanter hooks, with multiple setae on Setae VI. ABL = 23.0 to 30.0 mm; TW = 10.0 to 12.0 mm (from Gidaspow 1963).

FIGURE 7.—Habitus of Calosoma sayi Dejean, dorsal aspect, male from Cayuga, Guatemala.

FIGURES 8, 9.—Right middle tibia, male, lateral aspect: 8, C. angulatum Chevrolat, San Jose, Costa Rica; 9, C. fulgens Chaudoir, Posorja, Ecuador.

GEOGRAPHICAL DISTRIBUTION (Figure 20).—The range of this species extends from northern Argentina to Mexico.

NATURAL HISTORY.—According to Gidaspow (1963), these beetles have a broad habitat range, including dense lowland forests, deciduous scrub forest, grasslands, and cultivated fields throughout the tropical and subtropical regions of its distribution. In Central America, they occur in both TROPICAL DRY and MOIST FOREST. All specimens I studied were fully winged; the species is a highly vagile dispersant.

TAXONOMY.—Gidaspow (1963) discussed at length the taxonomic problems involved with this species and its sister species, C. sayi. I agree with her conclusion that there are two species involved and her application of names is correct; however I do not believe that subspecies are warranted given the present level of study on the material at hand; thus I regard C. granulation a junior synonym of C. alternans. Various other names have been associated with alternans at the subspecific level or less. It appears to me all these are based on a few specimens from different places and that no one has had good enough series to show separately evolving populations. Until this kind of study is done, I prefer to regard the species as monotypic, treating the various proposed names as synonyms.

MATERIAL EXAMINED.—Types (see above) and 10 specimens (Table 3).

07.01.002 Calosoma sayi Dejean

FIGURES 7, 12, 21

Calosoma sayi Dejean, 1826:198. [Lectotype female, “Amérique septenlrionale” {Say) (MNHP), herein selected.]

Calosoma armata LaPorte, 1835:156. [Lectotype male, MEXICO (Gory) (MNHP), herein selected.]

Calosoma sayi abdominale Géhin, 1885:58. [Lectotype male, MEXICO (MNHP), herein selected.]

Calosoma sayi virginica Casey, 1897:344. [Lectotype male, USA, Virginia, Norfolk (Casey) (USNM), herein selected.]

Calosoma cuprascens Roeschke, 1900:71. [Holotype male, probably Greater Antilles (depository unknown).]

DIAGNOSTIC COMBINATION.—These beetles are very similar to the preceding species except the elytral costae are more or less equally wide and only moderately raised. The males have two basal tarsal articles of the anterior leg clothed beneath with spongy pubescence. Color and luster: blackish with cupreous or greenish reflections; blackish to green on head and pronotum, mouthparts, appendages, and venter dark piceous. Form: size medium, narrow, females not wider than males; eyes large and prominent Structure: elytral intervals moderately convex, 3, 7, and 11 catenate with large, deep foveae, all transversely scaly; head rugosely punctate. Microsculpture finely isodiametric or on pronotum slightly transverse. Male with 2 dilated basal tarsal articles on foreleg, each with setiferous pad beneath; middle tibia with large brush of setae apicomedially; metatrochanter ventrally hooked apically; and male median lobe moderately bent laterally near apex. Female without tarsal pads, trochanter hooks; with multiple setae on sternum VI. ABL = 22.0 mm to 30.0 mm; TW = 9.0 to 12.0 mm (from Gidaspow, 1959).

FIGURES 10–19.—Pronotum. dorsal aspect: 10, C. fulgens Chaudoir, Posorja, Ecuador, 11, C. alternant (Fabricius), Alios de Maje, Panamá; 12, C. sayi Dejean, San Pedro Montes de Oca, Costa Rica; 13, C. abbreviation Chaudoir, Piura, Peru; 14, C. scrutator (Fabriciui), Nevada, Vernon County, Missouri; 15, C. aurocinctum Chaudoir, El Union, Tamaulipas, Mexico; 16, C. angulation Chevrolat, San Jose, Costa Rica; 17, C. marginalis Casey, Clinton, Custer County, Oklahoma; 18, C. glabratum Dejean, Bogota, Colombia; 19, C. ampliator Bates, Villa Lerdo, Durango, Mexico.

FIGURE 20.—Geographical distribution map of Calosoma alternans (Fabricius) in Central America.

GEOGRAPHICAL DISTRIBUTION (Figure 21).—The range of this species extends from the northeastern United States south to Panama and throughout the Greater Antilles.

NATURAL HISTORY.—In Central America, these beetles are found in TROPICAL DRY and VERY DRY FORESTS, TROPICAL MOIST FOREST, PREMONTANE MOIST, and WET FORESTS in cultivated lands, along streams and rivers, and in scrub lands, from sea level to 1210 m. All specimens studied are fully winged; the species is a highly vagile dispersant. They may be found in January, February, April, May, June, July, and August.

TAXONOMY.—Gidaspow (1963) discusses at length the taxonomic problems involved with this species and its sister species, C. alternans. I discuss the problem under that species (see above). I do not believe that subspecies are warranted given the present level of study on the material at hand; thus I regard C. abdominale Géhin and C. virginica Casey as junior synonyms of C. sayi. It appears to me all these are based on a few specimens from different places and that no one has had good enough series to show separately evolving populations. Until this kind of study is done, I prefer to regard the species as monotypic, treating all the various proposed names as synonyms.

MATERIAL EXAMINED.—Types (see above) and 47 specimens (Table 4).

TABLE 3.—Central American locality data deduced from specimen labels; 07.01.001 Calosoma alternans (Fabricius); see Figure 20.

Calosoma abbreviatum Chaudoir

FIGURE 13

Calosoma abbreviatum Chaudoir, 1869:371. [Holotype male, PERU or BOLIVIA (LaFerté-Sénectèrt) (MNHP).]

DIAGNOSTIC COMBINATION.—Among our species with foveate elytral intervals 3, 7, and 11, this species is easily recognized by its metallic green color, non-contrasting elytral foveae, and blunt metatrochanters. Color and luster: brassy green, forebody more green; mouthparts, appendages, and venter dark piceous. Form: size small to medium, relatively broad; female elytra expanded; eyes large and prominent Structure: elytral intervals shallowly convex, 3, 7, and 11 catenate with large foveae, all transversely scaly; head finely rugosely punctate. Microsculpture finely isodiametric, effaced from pronotum, pronotum with finely and coarsely punctate surface. Male with 3 dilated basal tarsal articles on foreleg, each with setiferous pad beneath; middle tibia with large brush of setae apicomedially; metatrochanter apically truncate; and male median lobe apex straight and broadly rounded. Female without tarsal pads; with blunt metatrochanter; and with multiple setae on sternum VI. ABL = 22.0 to 25.0 mm; TW = 8.5 to 10.0 mm (from Gidaspow 1963).

GEOGRAPHICAL DISTRIBUTION.—The range of this species extends from about 15°S to 3°N latitude along the west coast of South America. It can be expected to occur on the Azuero Peninsula or in the Darièn (when the latter is cut over for agriculture.)

NATURAL HISTORY.—According to Gidaspow (1963), these beetles live in cotton-growing regions of Peru, on the brush-covered plains of Colombia, and in sandy areas between the coast and oak covered foothills of Ecuador and Peru. Should the west coast of Panama (Darien) be opened to agriculture, it is possible these beetles could expand their range northward. These beetles are fully winged and no doubt fly quite well.

MATERIAL EXAMINED.—Type (see above) and 4 specimens from Peru.

07.01.003 Calosoma scrutator (Fabricius)

FIGURE 14

Carabus scrutator Fabricius, 1775:239. [Holotype, sex unknown, USA, Virginia (Banks) (BMNH).]

DIAGNOSTIC COMBINATION.—These large, metallic blue and green beetles with broad, inflated elytra are easily recognized on color alone, differing from members of C. aurocinctum by the former’s bicolored pronotum. Color and luster: black, with golden green spots near eyes, pronotum with dark blue or black disc and golden green or purplish margins, elytra metallic or dark green, venter with blue luster, appendages brown with blue luster. Form: size large, head and pronotum narrow relative to broadly inflated elytra; eyes large and prominent Structure: pronotum with sides evenly arcuate, small rounded hind angles, without basal setae; elytra serrate near humerus, with striatopunctate intemeurs, punctures connected by scaly cross lines, intervals convex; middle tibia markedly arcuate, in male with dense brush of reddish setae; male foretarsi with four dilated articles, each with spongy pubescence beneath. ABL = 25.0 to 35.0 mm; TW = 11.0 to 16.0 mm (from Gidaspow, 1959).

GEOGRAPHICAL DISTRIBUTION.—Although I have not seen any specimens, Gidaspow (1959, 1963) indicates that these beetles were found in Guatemala and Venezuela. They are relatively common also in Mexico.

FIGURE 21.—Geographical distribution map of Catatonia sayi Dejean in Central America.

NATURAL HISTORY.—Lindroth (1960) indicates that these beetles are found in hardwood forests where they feed on tent and noctuid caterpillars. They are fully winged and fly to lights at night.

MATERIAL EXAMINED.—Type (see above) and 20 specimens from the eastern United States.

07.01.004 Calosoma aurocinctum Chaudoir

FIGURE 15

Calosoma splendidum Perbosc, 1839:261. [Holotype female, MEXICO, Santa Domingo (MNHP).]

Calosoma aurocinctum Chaudoir, 1850:420. [New name for C. splendidum Perbosc, 1839:261, not Dejean.]

DIAGNOSTIC COMBINATION.—Pronotum without basal seta at hind angle; venter brilliant metallic, pronotum bluish green, not contrasting with elytra. Color and luster: bluish green, elytra often dark green with purple or golden margin, femur blue, tibiae and tarsi black. Form: size medium large, head and pronotum narrow relative to broadly inflated elytra; eyes large and prominent Structure: pronotum with sides evenly and markedly arcuate, small pointed hind angles, without basal setae; elytra serrate or not near humerus, with striatopunctate interneurs, punctures connected by shallow scaly cross lines, intervals convex; middle tibia markedly arcuate in male, slightly arcuate in female; male foretarsi with four dilated articles, each with spongy pubescence beneath. ABL = 22.0 to 30.0 mm; 10.0 to 14.0 mm (from Gidaspow, 1959).

TABLE 4.—Central American locality data deduced from specimen labels; 07.01.002 Calosoma sayi Dejean; see Figure 21.

GEOGRAPHICAL DISTRIBUTION.—Although Gidaspow (1963) reported this species from Nicaragua, I have seen no specimens from there and she does not give, in her paper, a list of depositories associated with each species. As several southern Mexican records are available, I have no doubt that this species could inhabit Central America.

NATURAL HISTORY.—Unknown.

MATERIAL EXAMINED.—Type (see above) and 10 specimens from Mexico.

07.01.005 Calosoma angulatum Chevrolat

FIGURES 9, 16, 22

Calosoma angulation Chevrolat, 1834, fasc. 2:44. [Holotype male, MEXICO, Vera Cruz, Boca del Monte (Lesueur) (MNHP).]

Calosoma angulicolle Chaudoir, 1869:377. [Lectotype female, COLOMBIA, Santa Marta (Fontanitr) (MNHP), herein selected.] [See also Breuning, 1928a: 103].

Calosoma uniforme Géhin, 1885:63. [Type, sex unknown, MEXICO, Mazatlan (MNHP).] [See also Breuning, 1928a: 101.]

Calosoma forrtri Géhin, 1885:64. [Type, male, ARIZONA (MNHP).] [See also Jeannel, 1940:203.]

DIAGNOSTIC COMBINATION.—Middle tibia straight (as in Figure 8); pronotum with angulate lateral margins, basal seta absent; elytral intervals convex, intemeurs finely punctate, punctures of adjacent interneurs connected with transverse grooves especially basally causing a markedly scaly appearance. Color and luster: black, base of pronotum and elytral margin with bluish or metallic green luster, venter black, shiny. Form: size medium large, elongate and narrow, head and pronotum narrow relative to broadly inflated elytra; eyes large and prominent Structure: pronotum with sides evenly and markedly arcuate, small pointed hind angles, without basal setae; elytra serrate near humerus, with deeply striatopunctate interneurs, punctures connected by scaly cross lines, intervals convex; middle tibia not arcuate in males or females; male foretarsi with three dilated articles, each with spongy pubescence beneath. ABL = 25.0 to 33.0 mm; TW = 9.0 to 12.0 mm (from Gidaspow, 1959).

GEOGRAPHICAL DISTRIBUTION (Figure 22).—Gidaspow (1959, 1963) indicates that this so called polytypic species has a range extending from the southern United States from Arizona to Texas to Colombia and Venezuela. I do not believe that subspecies are warranted given the present level of study on the material at hand especially from Central America, thus I regard C. angulatum Chaudoir and C. angulicolle Chaudoir as synonyms. It appears to me all these are based on a few specimens from different places and that no one has had good enough series to show separately evolving populations. Until this kind of study is done, I prefer to regard the species as monotypic.

NATURAL HISTORY.—These beetles have been collected from 280 to 317 m elevation and from 1000 to 1300 m elevation in PREMONTANE MOIST FOREST in Costa Rica and are often found flying around lights at night It is likely that they occur in cultivated land as well. They have been found in May through October.

MATERIAL EXAMINED.—Types (see above) and 12 specimens (Table 5).

07.01.006 Calosoma marginalis Casey

FIGURE 17

Calosoma lugubre LeConte, 1853:400. [Holotype male, TEXAS, Braunfels (Lindheimer) (MCZ).]

Calosoma marginalis Casey, 1897:340. [Holotype male, ?ARIZONA (Levette) (USNM).]

Calosoma leconlei Csiki, 1927:21. [New name for Calosoma lugubre LeConte, 1853:400; not C. lugubre Motschulsky.]

DIAGNOSTIC COMBINATION.—Middle tibia straight (as in Figure 8); pronotum with angulate lateral margins, basal seta absent; elytral intervals flat, intemeurs shallowly striatopunctate; metatrochanter asetose; head coarsely punctate. Color and luster: black, shiny, often with green luster on head, sides of pronotum, and elytral margin and foveae. Form: size large, head and pronotum narrow relative to broadly inflated elytra; eyes large and prominent Structure: pronotum with sides obtusely angulate, small triangular hind angles, without basal setae; elytra serrate near humerus, with fine striatopunctate intemeurs, punctures connected by shallow scaly cross lines, intervals flat; middle tibia nearly straight in male and female; male foretarsi with four dilated articles, each with spongy pubescence beneath. ABL = 23.0 to 32.0 mm; TW = 9.5 to 13 mm (from Gidaspow 1959).

GEOGRAPHICAL DISTRIBUTION.—Gidaspow (1963) indicated she saw two specimens from Costa Rica. However, I could not locate these specimens, and I have seen no other records of this species from Central America. Otherwise the species has a known range extending from Colorado south to Cucrnavaca, Mexico.

NATURAL HISTORY.—Members of this species can be found near wet or damp areas with shrubby vegetation or small trees in the high deserts of the American southwest and Mexico; it is likely they have become adapted to the margins of cultivated lands as well. They often fly to lights at night.

MATERIAL EXAMINED.—Types (see above) and 30 specimens from the United States and Mexico.

07.01.007 Calosoma glabratum Dejean

FIGURE 18

Calosoma glabratum Dejean, 1831:565. [Lectotype male, COLOMBIA (Goudot) (MNHP), herein selected.]

Calosoma bolivianum Géhin, 1885:65. [Type, male, “Bolivia” as originally given by Gelun (MNHP).]

FIGURE 22.—Geographical distribution map of Calosoma angulation Chcvrolat in Central America.

DIAGNOSTIC COMBINATION.—Middle tibia straight (as in Figure 8); pronotum with angulate lateral margins, angles rounded, basal seta absent; elytral intervals flat, intemeurs shallowly striatopunctate; metatrochanter unisetose, round at tip; head finely and densely punctate. Color and luster: black, venter dark brown. Form: size small, head and pronotum narrow relative to broadly inflated elytra; eyes large and prominent. Structure: pronotum with sides obtusely angulate, small pointed hind angles, without basal setae; elytra serrate near humerus, with fine striatopunctate intemeurs, punctures connected by shallow scaly cross lines at base, intervals flat; middle tibia straight in male and female; male foretarsi with three dilated articles, each with spongy pubescence beneath. ABL = 18.5 mm; TW = 8.0 to 10.5 mm (from Gidaspow, 1963).

GEOGRAPHICAL DISTRIBUTION.—Gidaspow (1963) indicated that she saw specimens from Panama; however, I could not locate these specimens, and I have seen no other records of this species from Central America. Otherwise, the species has a known range extending from Bolivia north to Colombia.

NATURAL HISTORY.—Members of this species are found in cultivated lands and in natural grasslands. They are fully winged and probably are strong flyers as are other members of the genus.

MATERIAL EXAMINED.—Types (see above) and 4 specimens from Colombia.

TABLE 5.—Central American locality data deduced from specimen labels; 07.01.005 Calasoma angulation Chevrolat; see Figure 22.

07.01.008 Calosoma ampliator Bates

FIGURE 19

Calosoma ampliatorBates, 1891:223. [Lectotype female, MEXICO, Durango, Villa Lerdo (Hoge) (BMNH), herein selected.]

DIAGNOSTIC COMBINATION.—Pronotum without basal seta at hind angle; venter black or piceous, no metallic reflections. Color and luster: black, venter black or dark brown, shiny. Form: size medium large, head and pronotum narrow relative to broadly inflated elytra; eyes large and prominent Structure: pronotum with sides markedly arcuate, small rounded hind angles, without basal setae; elytra serrate near humerus, with fine striatopunctate intemeurs, punctures connected by shallow scaly cross lines, intervals flat or slightly convex; middle tibia straight in male and female; male foretarsi with three dilated articles, each with spongy pubescence beneath. ABL = 21.0 to 27.0 mm; 8.5 to 11.0 mm (from Gidaspow, 1959).

GEOGRAPHICAL DISTRIBUTION.—Gidaspow (1959) indicates she saw specimens from Chiriquí, Panama; however, I could not locate these specimens, and I have seen no other records of this species from Central America. Otherwise the species has a known range extending from Chihuahua to Veracruz in Mexico.

NATURAL HISTORY.—Although unknown for sure, members of this species likely can be found near wet or damp areas with shrubby vegetation or small trees in the Chihuahuan Desert of Mexico; it is possible they will or have become adapted to the margins of cultivated lands as have other species in the genus.

MATERIAL EXAMINED.—Type (see above) and 11 specimens from Mexico.

Literature Cited

Ball, G.E.

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Ball, G.E., and T.L. Erwin

1969. A Synopsis of the Tribe Loricerini (Coleoptera: Carabidae). Canadian Journal of Zoology, 47:877–907.

Bates, H.W.

1881. Insecta, Coleoptera, Cicindelidae, Carabidae. In F.D. Godman and O. Salvin, editors, Biologia Centrali-Americana, 1:1–40. London: Taylor and Francis.

1891. Additions to the Carabideous Fauna of México, with Remarks on Some of the Species Previously Recorded. Transactions of the Entomological Society of London, 1891:223–278.

Breuning, S.

1927. Monographic der Gattung Calosoma Web. (Carab.). Koleopterologische Rundschau, 13:129–232.

1928a. Monographic der Gattung Calosoma Web. (Carab.). Weiner Entomologische Zeitung, 44:81–141.

1928b. Monographic der Gattung Calosoma Web. Koteopterologische Rundschau, 14:43–101.

1928c. Monographic der Gattung Ceroglossus Sol. (Col. Carab.). Eos, 4:283–338.

1931. Cinq nouvelles formes de Carabini. Bulletin du Muséum d'Histoire Naturelle, 3:620–623.

Casey, T.L.

1897. Coleopterological Notices, VII Annals of the New York Academy cf Science, 9:285–684.

1913. Studies in the Cicindelidae and Carabidae of America. Memoirs on the Coleoptera, 4:1–192.

Chaudoir, M. de

1850. Memoire sur la famille des Carabiques. Bulletin de la Société Impériale des Naluralistes de Moscou, 23:349–460.

1863. Description de cicindélétes et de Carabiques nouveaux. Revue et Magasin de Zoologie, 15:111–120.

1869. Descriptions de Calosoma nouveaux des collections de MM de Chaudoir et Salle. Annates de la Société Entomologique de France, 9:367–378.

Chevrolat, L.A.A.

1834. Coléoptères du Mexique. Fascicle 2, 44 pages. Strasbourg.

Csiki, E.

1927. Carabinae. Coleopterorum catalogus, Carabidae, pars 91, 1:1–314.

Darlington, P.J., Jr.

1952-1971. The Carabid Beetles of New Guinea, Parts 1-2. Bulletin of the Museum of Comparative Zoology, parts 1-4. [1952, Part 2, The Agonini, 107(3):89–252, 4 plates; 1962, Part 1, Harpalinae through Pterostichini, 126(3):321–564, 4 plates; 1968, Part 3, Harpalinae (Continued): Perigonini to Pseudomorphini, 137(l):l-253; 1971, Part 4, General Considerations, Analyses and History of Fauna, Taxonomic Supplement. 142(2):130–337.]

1957. Zoogeography: The Geographical Distribution of Animals. 675 pages. New York: John Wiley and Sons, Inc.

Dejean, P.F.M.S.

1826. Spécies général des Coléoptères de la collection de M. de Comte Dejean. Volume 2, 501 pages. Paris.

1831. Spécies général des Coléoptères de la collection de M. de Comte Dejean. Volume 5, 883 pages. Paris.

Duméril, A.M.C.

1806. Zoologie analytique, ou méthode naturelle de classification des animaux. 343 pages. Paris.

Erwin, L.J.M.

1976. Application of a Computerized General Purpose Information Management System (SELGEM) to a Natural History Research Data Bank (Coleoptera: Carabidae). Coleopterist Bulletin, 30(1):1-32.

Erwin, T.L.

1970. A Reclassification of Bombardier Beetles and a Taxonomic Revision of the North and Middle American Species (Carabidae: Brachinida). Quaestiones Entomologicae, 6:4–215.

1973. A Supplement to the Bombardier Beetles of North and Middle America: New Records for Middle America (Coleoptera: Carabidae). Coleopterist Bulletin, 27:79–82.

1974. Studies of the Subtribe Tachyina (Coleoptera: Carabidae: Bembidiini). Part II: A Revision of the New World-Australian Genus Pericompsus LeConte. Smithsonian Contributions to Zoology, 162: 96 pages.

1975. Studies of the Subtribe Tachyina (Coleoptera: Carabidae: Bembidiini). Part III: Systematics, Phylogeny, and Zoogeography of the Genus Tachyta Kirby. Smithsonian Contributions to Zoology, 208: 68 pages.

1978. Techniques. Coleopterist Bulletin, 32(4):372.

1979. Thoughts on the Evolutionary History of Ground Beetles: Hypothesis Generated from Comparative Faunal Analysis of Lowland Forest Sites in Temperate and Tropical Regions. In T.L Erwin, G.E. Ball, D.R Whitehead, and A.L. Halpem, Carabid Beedes: Their Evolution, Natural History, and Classification. In Proceedings of the First International Symposium of Carabidology, pages 539-592. The Hague: Dr. W. Junk.

1982a. Small Terrestrial Ground Beetles of Central America (Carabidae: Bembidiina and Anillina). Proceedings of the California Academy of Science, 42(I9):455–496.

1982b. Agra, Arboreal Beedes of Neotropical Forests: erythropus Group Systematics (Carabidae). Systematic Entomology, 7:39–71.

1985. The Taxon Pulse: A General Pattern of Lineage Radiation and Extinction among Carabid Beedes. In G.E. Ball, Taxonomy, Phylogeny, and Zoogeography of Beetles and Ants: A Volume Dedicated to the Memory of Philip Jackson Darlington Jr. 1904-1953, pages 437-472. The Hague: Dr. W. Junk.

Erwin, T.L., G.E Ball, D.R. Whitehead, and A.L Halpem, editors

1979. Carabid Beetles: Their Evolution, Natural History, and Classification. In Proceedings of the First International Symposium of Carabidology, 635 pages. The Hague: Dr. W. Junk.

Erwin, T.L., and D.K. Kavanaugh

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., and LL. Sims

1984. Carabid Beetles of the West Indies (A Generic Synopsis). Quaestiones Entomologicae, 20:351–464.

Fabricius, J.C.

1775. Systema Entomologiae. 832 pages. Lipsiae [Leipzig).

1792. Entomologia Systematica. 538 pages. Hafniae [Copenhagen].

Géhin, J.J.B.

1885. Catalogue synonymique el systematique des Coléoptères de la tribu Carabides. 104 pages. Remiremont.

Gidaspow, T.

1959. North American Caterpillar Hunters of the Genera Calosoma and Callisthenes (Coleoptera, Carabidae). Bulletin of the American Museum of Natural History, 110:229–343.

1963. The Genus Calosoma in Central America, the Antilles, and South America (Coleoptera, Carabidae). Bulletin of the American Museum of Natural History. 124:279–313.

Jeannel, R.

1940. Les Calosomes. Mémoires du Muséum National d'Histoire Natu-rellede Paris, 13:1–240.

Kavanaugh, D.K., and J. Negre

1983. Notiokasiini—A New Tribe of Carabidae (Coleoptera) from Southeastern South America. Coleopterist Bulletin, 36(4):549–566.

Kirby. W.

1837. Insects: Coleoptera. In Sir J. Richardson, Fauna Boreali-Americana; or the Zoology of the Northern Parts of British America, 4: 249 pages, 8 plates. Norwich.

Laporte, F.L.N.C.C. de

1835. Études entomologiques, ou descriptions d'insects nouveaux, et observations sur la synonymie. Volume 1, pages 95-159. Paris.

Lapouge, G.V. de

1924. Calosomes nouveaux ou mal connus (Col. Carabidae). Miscellanea Entomologies, 28:37–44.

Latreille, P.A.

1802. Hisioire naturelle, générale et particulière des crustaces et des insectes. Volume 3, 467 pages. Paris.

1810. Considérations générales sur l'ordre naturel del animaux composant les classes des crustacés, des archnides, et des insectes; avec un tableau méthodique de lews genres, disposé en families. 444 pages. Paris.

LeConte. J.L.

1853. Notes on the Classifications of the Carabidae of the United States. Transactions of the American Philosophical Society, 10:363–403.

Lindroth, C.H.

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Motschulsky, V. von

1865. Enumération des nouvelles espèces de coléoptères rapportés de ses vogages. Bulletin de la Sociélé Impériale des Naturalistes de Moscou, 38:227–313.

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Reichardt, H.

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Reiche, L.

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Roeschke, H.

1900. Carabologische Notizen, VI. Enlomologische Nachrichten, 26:68–72.

Weber, F.

1801. Observations entomologicae, continentes novoroum quae condidit generum characteres, el nuper detectorum specierun descriptiones. 116 pages. Kiliae.

Westwood.J.O.

1840. Synopsis of the Genera of British Insects. In An Introduction to Modern Classification of Insects; Founded on the Natural Habits and Corresponding Organization of the Different Families, supplement, volume 2,587 pages. London: Longman, Orme, Brown, Green and Longman.

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