CHARLES A. TRIPLEHO A Review of the Qenus

_ Zopherus of the World

(Coleoptera: i

Tenebrionidae)

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 108

Charles A. Triplehorn A Revie w o f th e

Genus Zopherus

of the World

(Coleoptera:

Tenebrionidae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

U)72
ERRATA FOR SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY,

NUMBER 108

Third line from bottom in Abstract should read:

valid. Three new species are described: Z. solieri (Mexico), Z. xestus (Texas), and

Page 6, item 4 should read:

4. Dorsum of both pronotum and elytra largely white, especially laterally.

nodulosus haldemani Horn

Dorsum of both pronotum and elytra largely black; black spots not confined to medial

areas (3) nodulosus nodulosus Solier*
ABSTRACT

Triplehorn, Charles A. A Review of the Genus Zopherus of the World (Coleoptera:

Tenebrionidae). Smithsonian Contributions to Zoology, number 108, 24 pages, 4

plates. 1972. — The genus Zopherus consists of 19 species and 2 subspecies which inhabit the Western Hemisphere from western North America to Venezuela. The three

Casey genera, Megazopherus, Zopherinus, and Zopherodes, are placed in synonymy

with Zopherus, and of the 23 names proposed by Casey, only two are considered

valid. Three new species are described: Z. solieri (Mexico), Z. Xestus (Texas), and

Z. championi (Mexico and Texas). A key to the known species of Zopherus and

a discussion of each species is presented.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

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Stock Number 4700-0169
Charles A. Triplehorn A Review of the

Genus Zopherus

of the World

(Coleoptera:

Tenebrionidae)

Introduction

The most recent comprehensive survey of this interesting group of beetles was by Casey (1907a). In his

paper he divided the genus Zopherus into four genera, three of which (Megazopherus, Zopherinus, and

Zopherodes) were new, and he described a number

of new species which were included in keys along with

his concept of previously known species. It should

be pointed out that Casey did not always correctly

establish the identity of species described by earlier

workers and consequently made many errors and

created numerous synonyms.

In the same year Casey (1907b) described eight

more new species of Zopherodes which were, as was

his custom, presented in the form of a key. This key

did not include previously described species, so the

reader was forced to use two separate keys. In my

work with the Tenebrionidae, I found it impossible

to use either key. As it turned out, all of the species

described in the second paper (1907b) are synonyms

of previously described species, but until this was established, determinations at the species level were

uncertain.

Blaisdell (1931) presented a key to the California

species of Zopherodes from which he omitted Z. graniCharles A. Triplehorn, Department of Entomology, The

Ohio State University, Columbus, Ohio 43210.

collis Horn, perhaps the most characteristic member

of the genus in the California fauna. He also described

Z. sanctaehelenae in that paper.

The purpose of the present paper is to present a

key to the known species of Zopherus, to clarify a

considerable number of synonyms, and to summarize

our knowledge to date of the biology and distribution

of the group.

History

Gray (1832) published brief descriptions with colored

illustrations of two new species of Coleoptera which

established the starting point for the history of the

genus Zopherus. He identified one of these (plate

124: figure 3) in the caption (page 796) as "Zophorus chilensis, the type of a new genus near Zophosis." The other species (plate 50: figure 5) he called

Zophorus mexicanus.

Laporte (1840: 205) called attention to an error

in translitteration from the Greek and spelled the

name "Zopherus." It is clear that he was only emending Gray's spelling and therefore should not be cited

as the author of the genus as many subsequent authors and catalogers have done.

Solier (1841:30) placed Zopherus in his tribe Zopherites, along with Nosoderma Dejean and Diceroderes Solier. It is interesting that he cited Hope as
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

author of Zopherus and listed nervosus, nodulosus,

mexicanus, and laevicollis, all of which are now valid

species. No mention is made of chilensis.

Hope (1840:110) likewise is often cited as the

author of Zopherus and of Zopherus mexicanus. He

merely mentioned the generic and specific name and

the locality (Mexico). He cannot possibly be credited

with authorship of the genus since, at best, his name

would be a nomen nudem.

Lacordaire (1859:91) followed Solier in the tribal

arrangement, his spelling of the tribe differing slightly (Zopherides). He also correctly cites Gray as the

author of Zopherus.

Zopherus was the genus into which LeConte, Horn,

and Champion placed the species which they described. The genus was thus well known and very

stable until Casey (1907a) divided it into four genera. He designated a type species for each as follows:

Megazopherus containing only chilenis Gray; Zopherodes with tristis LeConte as type species; Zopherinus with limbatus Casey as type species, and Zopherus Gray with mexicanus Gray as type species.

In the same year Casey (1907b) published a larger

paper on North American Tentyriinae in which he

assigned the four genera listed above to the tribe

Zopherini. Gebien (1910:93) classified the group in

the subfamily Zopherinae which included essentially

Casey's tribes Zopherini and Nosodermini.

Boving and Craighead (1931:41), basing their conclusions for the most part on larval studies, created

a separate family, Zopheridae, for those beetles in

Casey's tribes Zopherini and Nosodermini. Family

status for the group was also given by Crowson

(1955:116, 127), Boddy (1965:77), and Watt (1967:

82). I do not feel that there are strong enough arguments for a separate family and I concur with Arnett

(1963:662), who places the group in the tribe Zopherini, subfamily Asidinae.

Morphology

All members of the genus Zopherus are elongate and

subcylindrical, with extremely thick and hard integuments. In pinning them it is frequently necessary to

first drill a hole in the elytra before the pin can be

inserted. Despite this hardness, the elytra and pronotum are easily scored with a sharp instrument, the

integument having the physical properties of an extremely hard wax.

COLORATION.—North of the Rio Grande River

all but two of the species are entirely black or have

faint traces of white or gray on the flanks of the

pronotum. In the New World Tropics, all of the

species are patterned, sometimes boldly, in black and

white. Although these colors provide useful characters,

they are subject to considerable variation, much of it

apparently due to abrasion. Moreover, soon after

death, the body rapidly exudes grease which hardens

to a tarlike consistency, causing even the boldly patterned black and white species to become uniformly

shiny black. Surface sculptures (punctures and tubercles) are likewise obliterated by the grease. It is

necessary, therefore, to soak specimens in ether or

other grease solvent for at least 30 minutes before

drawing conclusions in regard to color and sculpture.

I have repeatedly (up to eight times) restored the

color in a number of specimens in this manner, only

to find them completely black again in less than a

week. Luster ranges from shiny to dull.

SCULPTURE.—Most of the species of Zopherus

have tubercles of some sort on the elytra. These may

be in the form of large, slightly raised areas or they

may be small and distinctly verrucose. The pronotum

may be tuberculate, granulate, or simply punctured,

and this is also true of the prosternum, prosternal

process, and ventral surface of the pronotum. Surface

sculpture provides many of the most useful characters

for species discrimination, even though there is much

intraspecific variation.

HEAD. — The head is retracted into the prothorax

beyond the eyes, which are thus entirely concealed

from above when the head is in repose. Blaisdell

(1931:113) was the first to point out the enormous

size of the clypeus, which occupies three fourths or

more of the exposed front of the head. The feebly indicated frontal sutures extend backward and inward

to just before the eyes and join on the interocular

surface, which is also concealed from above when

the head is in repose. The labrum is transverse and

its anterior margin is densly fringed with rather long,

golden hairs. The mandibles are massive, broadly

truncate, or feebly bidentate apically. The eyes are

narrow, transverse, flattened, and very finely facetted.

ANTENNAE. — The antennae are short and stout,

with the apical three segments fused into a compact,

oblong club. An elongate patch of yellow hairs is

present at the apex of the club, with smaller patches

of yellow hairs on the apicolateral portions of seg-
NUMBER 10 8

ments 9 and 10 (Figure 7). The antennae are usually

retained within deep, entire fossae located on the

ventral side of the prothorax of all species. These

fossae are usually rounded apically (Figure 6) but

in some species they are appendiculate (Figure 5).

PRONOTUM. — The pronotum is usually longer than

broad, the apical margin is always broader than the

basal margin, and the lateral margins are frequently

prominent or even angulate opposite the apex of the

antennal fossae on the ventral surface. A distinct

marginal bead is never present laterally or apically

but the base may be reflexed.

SCUTELLUM. — The scutellum is small and inconspicuous, usually hidden beneath the base of the pronotum when the body is in repose.

ELYTRA. — The elytra are fused but the sutural line

is always apparent. The apices are always conspicuously sculptured, either with deep grooves or prominent,

flattened or globose tubercles. The flanks of the elytra

are rounded and the epipleura are absent.

ABDOMEN.—The terminal abdominal sternum presents some interesting modifications which are useful

taxonomically. These are folds or ridges, referred to

in this paper as "calli," and are discussed under each

species. Usually a conspicuous, compact patch of

golden hairs is present at the extreme apex of the

terminal abdominal sternum.

LEGS. — The ventral surfaces of the femora, tibiae,

and tarsi are clothed on each side with dense and extremely fine and compact ridges of golden hairs that

extend from base to apex of each extremity, and conspicuously outline the bare central portion. These

ridges of hair also occur around the periphery of the

distal end of the tibiae, and similar dense patches of

hair completely clothe the trochanters and form a

pointed tuft at the apex of each tarsus between the

claws.

SEXUAL CHARACTERS. — I have found no external

morphological characters to separate the sexes. Females tend to be somewhat larger and stouter than

males, but body proportions are, in general, unreliable in determining sex. Neither male nor female

genitalia proved useful in species discrimination. The

aedeagus of Z. chilensis is illustrated (Figure 8).

Classification

The three genera, Megazopherus, Zopherodes, and

Zopherinus, described by Casey are untenable, and it

is necessary to reduce them all to synonyms of Zopherus. That they fall into fairly well-defined species

groups cannot be denied, but to attempt to force

them into a rigid system of higher categories creates

more problems than it solves. Casey, as was his

practice, based the primary taxonomic category —

the species — one step lower than he should have.

Most of the forms he considered to be species are

herein shown to be merely normal intraspecific variations.

Zopherus is a perfect group to illustrate Casey's

approach to taxonomy. Of the 23 species described in

his three papers (1907a, 1907b, 1924), only two

names are retained — uteanus, a valid species, and

ventriosus, which is here considered a subspecies of

granicollis. Casey redescribed elegans Horn four times

and tristis LeConte three times. He created seven

synonyms of gracilis Horn, at the same time failing to

identify any of his specimens as gracilis. Sixteen of the

23 species were based on unique specimens, and it

soon became evident to me that Casey was describing individuals and not species. He felt obligated to

place all of the described species into his expanded

system, and he had to guess on the placement of a

number of them. On many of them he guessed wrong.

Synoptic Catalog of Zopherus

Zopherus Gray, 1832:796.

Megazopherus Casey, 1907a: 36.

Zopherinus Casey, 1907a:36.

Zopherodes Casey, 1907a: 36.

1. chilensis Gray, 1832: 796.

insignis Blanchard, 1861: pi. 7A, fig. 6.

bremei Guerin-Meneville, 1844:18.

moreletii Lucas, 1852:XXIII.

2. nervosus Solier, 1841:42.

pectoralis LeConte, 1851:130.

reticulatus (var.) Champion, 1884:43.

compactus Champion, 1884:43.

marmoratus Casey, 1907a: 36.

3. nodulosus nodulosus Solier, 1841:43.

variolosus Sturm, 1843:349.

sallaei (var.) Champion, 1884:42.

verrucosus (var.) Champion, 1884:43.

nodulosus haldemani Horn, 1870:271.

4. jourdani Salle, 1849:301.

mexicanus (auct., nee. Gray)

costaricensis Champion, 1884:40.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

5. jansoni Champion, 1884:39.

6. mexicanus Gray, 1832:796.

maculatus Champion, 1884:41.

7. angulicollis Champion, 1884:42.

8. laevicollis Solier, 1841:46.

venosus (var.) Champion, 1884:40.

tuberculatus Champion, 1884:41.

limbatus (Casey), 1907a:37.

9. xestus, new species

10. solieri, new species

elegans Champion (nee. Horn), 1892:489.

11. championi, new species

elegans Champion (nee. Horn), 1892:489.

12. tristis LeConte, 1851:130.

aequalis (Casey), 1907a:38.

variabilis (Casey), 1907b:464.

incrustans (Casey), 1907b:464.

13. concolor LeConte, 1851:130.

guttulatus Horn, 1867:160.

morosus (Casey), 1907b:465.

14. gracilis Horn, 1867:162.

pudens (Casey), 1907a:40.

caudalis (Casey), 1907a:41.

lugubris (Casey), 1907a:41.

pruddeni (Casey), 1907a:41.

luctuosus (Casey), 1907a:41.

elongatus (Casey), 1907b:466.

geminatus (Casey), 1907b:467.

15. uteanus (Casey), 1907a:40.

mormon (Casey), 1907a:40.

16. granicollis granicollis Horn, 1885:160.

induratus (Casey), 1907a:39.

californicus (Casey), 1907b:466.

prominens (Casey), 1924:305.

granicollis ventriosus (Casey), 1907a:39.

parvicollis (Casey), 1907b:466.

17. opacus Horn, 1867:161.

18. elegans Horn, 1870:272.

otiosus (Casey), 1907a:39.

verrucipennis (Casey), 1907b:465.

circumductus (Casey), 1924:304.

woodgatei (Casey), 1924:305.

19. sanctaehelenae (Blaisdell), 1931:111.

Acknowledgments

None of the species of Zopherus may be considered

common and some are very rare. The 2,429 specimens upon which this report is based came from

numerous sources, and I am indebted to many institutions and individuals for lending their collections to me and for allowing me to retain them for

an extended period.

I am especially grateful to Miss C. M. F. von

Hayek of the British Museum (Natural History) for

making available to me virtually all of the specimens

entrusted to her care, especially the George C. Champion material, which proved invaluable in working

out the Central American species. Through the courtesy of the late Dr. Harold J. Grant, Jr., of the

Academy of Natural Sciences of Philadelphia, I was

able to study all of Horn's material. Mr. T. J. Spilman

allowed me free use of the Casey collection at the

United States National Museum and helped in countless other ways. To these three I express a special

note of thanks.

A number of people have made a special effort to

collect specimens of Zopherus for me. Prominent

among these are Frank J. Moore, Josef N. Knull,

William Tyson, Richard L. Berry, my wife, Wanda

Elaine, and two sons, Bradley A. and Bruce W. Triplehorn. All of the photographs were taken by Richard

C. Arnold of the Photography Department, The Ohio

State University, whose painstaking skill with the

camera and interest in the project is evident. Donald

J. Borror and T. J. Spilman read the manuscript and

contributed many valuable suggestions. The efforts

of all these people are greatly appreciated.

Field work was partially supported by the American

Philosophical Society in 1963 (Grant No. 3091-Penrose Fund) and 1967 (Grant No. 4597-Penrose

Fund). Grateful acknowledgment is made to this society for its aid.

The following list of abbreviations represents the

deposition of specimens mentioned in this paper and

the individuals responsible for initiating the loans to

me:

AMNH American Museum of Natural History, New

York: Patricia Vaurie and Lee H. Herman, Jr.

ANSP Academy of Natural Sciences of Philadelphia: Harold J. Grant, Jr.

ASU Arizona State University, Tempe: Frank F.

Hasbrouck and Stanley C. Williams.

BMNH British Museum (Natural History), London: C. M. F. von Hayek and J. BalfourBrowne.
NUMBER 10 8

BYU Brigham Young University, Provo, Utah:

Vasco M. Tanner.

CAS California Academy of Sciences, San Francisco: Hugh B. Leech.

CAT Charles A. Triplehorn private collection.

CMP Carnegie Museum, Pittsburgh: George Wallace.

CNC Canadian National Collection, Ottawa:

Henry F. Howden and J. M. Campbell.

CU Cornell University, Ithaca, New York:

L. L. Pechuman and Henry Dietrich.

DEB David E. Bixler private collection.

DZSP Departamento de Zoologia, Sao Paulo, Brazil : F. S. Pereira and Hans Reichardt.

FMNH Field Museum of Natural History, Chicago,

Illinois: Rupert L. Wenzel.

FSCA Florida State Collection of Arthropods,

Gainesville: Robert E. Woodruff and

Howard V. Weems, Jr.

HFH Henry F. Howden private collection.

INHS Illinois Natural History Survey, Urbana:

Leonora K. Gloyd and Milton W. Sanderson.

INIA Instituto Nacional de Investigaciones Agricolas, Chapingo, Mexico: Francisco Pacheco M. and Alejandro Ortega C.

JAB Joseph A. Beatty private collection.

JMC J. M. Campbell private collection.

KSU Kansas State University, Manhattan: Carl

W. Rettenmeyer.

KWB Kirby W. Brown private collection.

LBSC Long Beach State College, Long Beach,

California: Elbert L. Sleeper.

MCZ Museum of Comparative Zoology, Harvard

University, Cambridge, Massachusetts:

John F. Lawrence and P. J. Darlington,

Jr.

MINN University of Minnesota, St. Paul: Edwin

F. Cook.

MSU Michigan State University, East Lansing:

Roland L. Fischer.

NC Neil Chernoff private collection.

OSU The Ohio State University, Columbus.

RLB Richard L. Berry private collection.

RS Ronald Somerby private collection.

TAM Texas Agricultural and Mechanical University, College Station: Horace R. Burke.

UA University of Arizona, Tuscon: Floyd G.

Werner.

UAlb University of Alberta, Edmonton: George

E. Ball.

UCB University of California at Berkeley: Paul

D. Hurd, Jr., and Jerry Powell.

UCD University of California at Davis: Robert O.

Schuster.

UCR University of California at Riverside: Everett I. Schlinger.

UIDA University of Idaho, Moscow: William F.

Barr.

UMo University of Missouri, Columbia: Wilbur

R. Enns.

UNA University of Northern Arizona, Flagstaff:

C. D. Johnson.

UNeb University of Nebraska, Lincoln: Warren T.

Atyeo.

UNev University of Nevada, Reno: Ira La Rivers.

USNM United States National Museum, Washington, D.C.: T. J. Spilman.

USU Utah State University, Logan: George F.

Knowlton.

WR William Rosenberg private collection.

WT William Tyson private collection.

VMK Vernon M. Kirk private collection.

Key to World Species of Zopherus

(*Species has not been taken in the United States)

1. Elytra abruptly inflexed at the sides, the inflexed sides more or less flat and having the

aspect of true epipleura; body feebly constricted between thorax and abdomen

(1) chilensis Gray*

Elytra not abruptly and less widely inflexed at sides; body strongly constricted between

thorax and abdomen 2

2. Elytra distinctly quadrituberculate at apex, the inner pair of tubercles smaller and more

terminal in position 3

Elytra bituberculate at apex or with only a short oblique ridge on each side of apex .. ... 5
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3. Ventral surface of pronotum with coarse tubercles (2) nervosus Solier*

Ventral surface of pronotum smooth 4

4. Dorsum of both pronotum and elytra largely white, especially laterally, nodulosus haldemani

Horn Dorsum of both pronotum and elytra largely black; black spots not confined to

medial areas (3) nodulosus nodulosus Solier*

5. Elytral apex distinctly bituberculate 6

Elytral apex with only oblique swollen ridges on each side 11

6. Apical elytral tubercles globose and prominent 7

Apical elytral tubercles large and flattened °

7. Base of elytra strongly reflexed, prominent (5) jansoni Champion*

Base of elytra rounded, not prominent (4) jourdani Salle*

8. Apical elytral tubercles almost contiguous at base, suture not swollen between them (Figure

2) ; color dull black (7) angulicollis Champion*

Apical elytral tubercles well separated at base with suture strongly swollen between them

(Figure 1) ; color either entirely black or black and while 9

9. Ventral callus of terminal abdominal sternum feebly arcuate at base (Figure 4)

(6) mexicanus Gray*

Ventral callus of terminal abdominal sternum strongly trilobed (Figure 3) 10

10. Dorsum more or less tuberculate or coarsely wrinkled; surface usually with considerable

amount of white (8) laevicollis Solier*

Dorsum almost smooth, usually entirely black (9) xestus, new species

11. Lateral portions of elytra and pronotum white 12

Entire dorsum unicolorous 14

12. Pronotum with punctures more or less muricate; prosternum with strong tubercles anterior

to coxae; apical elytral ridges prominent and separated from sutural ridge by a distinct

groove (18) elegans Horn

Pronotum with simple punctures; prosternum coarsely punctured, not tuberculate; apical

elytral ridges feebly developed and continuous (at least at base) with elevated sutural

ridge 13

13. Pronotum with lateral white borders having numerous black tubercles scattered over surface;

apical elytral ridges separated from raised sutural ridge except at extreme apex; lateral

areas of elytra more black than white; length 17-20 mm (10) solieri, new species*

Pronotum with lateral white borders having only a few minute flecks of black; apical

elytral ridges broadly in contact with raised sutural ridge; lateral areas of elytra more

white than black; length 12-15 mm (11) championi, new species

14. Elytra smooth or with minute, widely scattered muricate punctures; sometimes with scattered vermiculate lines (14) gracilis Horn

Elytra with distinct tubercles 15

15. Elytra abruptly narrowed at base and apex (humeri forming rather broad "shoulders") ;

elytra abruptly and steeply declivous behind and with margins parallel (12) tristis LeConte

Elytra gradually narrowed to base (humeri not forming distinct "shoulders") ; elytra

gradually declivous behind and with margins arcuate 16

16. Pronotal punctures simple, widely separated; elytra with large, prominent, more or less

contiguous tubercles (13) concolor LeConte

Pronotal punctures more or less muricate, sometimes granulate, at least laterally; elytral

tubercles variable, either large or fine 17

17. Elytral tubercles small, subequal in size to those of pronotum, widely separated 18

Elytral tubercles always much larger than those of pronotum, closely spaced, sometimes

contiguous 19

18. Dorsal surface of elytra between tubercles shiny (19) sanctaehelenae Blaisdell

Dorsal surface of elytra between tubercles dull, opaque . (17) opacus Horn

19. Clypeal punctures coarse, deep and closely spaced; robust species, pronotum usually narrower than elytra 20

Clypeal punctures small, shallow and widely separated; species of slender proportions,

pronotum usually as wide as or wider than elytra 21
NUMBE R 10 8

20. Elytral tubercles large, their bases usually more or less contiguous; lateral tubercles similar

to those of dorsum (16) granicollis granicollis Horn

Elytral tubercles smaller, well separated, lateral ones conspicuously transversely elongate

granicollis ventriosus (Casey)

21. Elytral tubercles subequal in size, uniformly and densely distributed (15) uteanus (Casey)

Elytral tubercles of two distinct sizes—large, shiny, conspicuous ones and small to minute

ones sparsely scattered over opaque areas between large ones.

(18) elegans Horn (melanistic phase)

1. Zopherus chilensis Gray

PLATE 2: FIGURE 9

Zophorus chilensis Gray, 1832:796, tab. 124, fig. 3.

Zopherus bremei Guerin-Meneville, 1844:18.—Breme, 1844:

307, tab. 9, fig. 2.

Zopherus Moreletii Lucas, 1852:xxiii (nomen nudem).

Zopherus insignis Blanchard, 1861: tab. 7a, fig. 6.

Zopherus chiliensis, Champion, 1884:38.

Zopherus chilensis, Champion, 1892:488.

Megazopherus chiliensis, Casey, 1907a: 36.

Megazopherus chilensis, Casey, 1907b: 461.

This, the largest species in the genus, may be recognized by the abruptly inflexed sides of the elytra which

resemble true epipleura. The constriction between

pronotum and elytra is less than in any other member

of the group. The size and coloration are extremely

variable, but most specimens show large amounts of

white. Usually the entire dorsum is a dull white with

a few dark tubercles showing through. The apex of

the elytra is distinctly bituberculate, each tubercle

globose and prominent.

I have seen none of the types, but there can be little

doubt that all of the names listed above refer to the

same species. Both Champion and Casey originally

spelled the specific name "chiliensis" and later spelled

it "chilensis," which is the correct spelling. Both

authors credited the species to Gray.

This insect has entered the popular literature under

the name "jewelled beetle" or "Makech." Jewels,

pearls, and bits of chenille are glued to the elytra and

pronotum, and the living beetle is tethered by a gold

chain as a living piece of costume jewelry. Many

specimens have been confiscated at United States

ports of entry, although it is doubtful that the insect

would become established, much less a pest of any

consequence.

MEASUREMENTS.—Length 34-46 mm; width 13—17

mm.

SPECIMENS EXAMINED.—-73 from the localities that

follow. COLOMBIA: Bonda, Papare. COSTA RICA:

Chiriqui Bay. EL SALVADOR: Quetzaltepeque, Santa

Tecla . GUATEMALA: Asuncion Mita , Jutiapa .

HONDURAS : Naco River near Corradia, San Pedro

Sula. MEXICO: Chilpancingo and La Union in

Guerrero; Merida and Progreso in Yucatan; 24 miles

northwest of Ocozocoautla in Chiapas; Xalostoc in

Morelos; Tehuantepec and Achazumba in Oaxaca.

PANAMA: Santa Fe in Darien. VENEZUELA: Puerto

Cabello. Specimens have been taken in every month of

the year.

2. Zopherus nervosus Solier

PLATE 2: FIGURE 10

Zopherus nervosus Solier, 1841:42, tab. 2, figs. 8-15.—

Champion, 1884:43; 1892:490.

Zopherus pectoralis LeConte, 1851:130.

Zopherus nervosus var. reticulatus Champion, 1884:43.

Zopherus compactus Champion, 1884:43, tab. 2, fig. 4 [new

synonymy].

Zopherus marmoratus Casey, 1907a:36 [new synonymy].

This distinctive species is largely black. Both the

thorax and elytra are strongly convex, and the ventral

surface of the pronotum is always coarsely and conspicuously tuberculate. The ventral callus of the apical

abdominal sternum is evenly and rather feebly

emarginate, not much swollen, and the associated

patch of yellow hairs is much smaller than is usual in

other species. In all specimens examined, the apex

of the antennal fossa is appendiculate.

The names listed above in synonymy appear to

represent merely color or slight structural variations of

the same species. I have studied 13 specimens of

Champion's Biologia Centrali-Americana series, representing those named by him as compactus, nervosus,

and variety reticulatus. In addition I have examined

Casey's 4 specimens of Z. marmoratus from Guerrero,

Mexico, and have labeled and hereby designate the

first specimen of the series as lectotype. All 4 specimens

also bear a USNM type number 46343.
8

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

All specimens I have seen range from very dark

(compactus) to mostly white (reticulatus, marmoratus), with typical nervosus intermediate between the

two extremes. The lightest specimens seen are from

Omilteme and Mochitlan, Guerrero.

MEASUREMENTS.—Length 17.0—30.5 mm; width

6.4-12.6 mm.

SPECIMENS EXAMINED.—35 from the localities that

follow. MEXICO: Mochitlan and Omilteme in Guerrero; La Parada, Juquila, Tepanzacoalco, and 6.6

miles south of Valle Nacional in Oaxaca; Jalapa and

Orizaba in Veracruz.

3. Zopherus nodulosus nodulosus Solier

PLATE 2: FIGURE 11

Zopherus nodulosus Solier, 1841:43.—Champion, 1884:42;

1892:489, tab. 22, fig. 6.

Zopherus variolosus Sturm, 1843:349, tab. 5, fig. 2.—Haldeman, 1852:376.

Zopherus nodulosus var. sallaei Champion, 1884:42;

1892:490.

Zopherus haldemanni var. verrucosus Champion, 1884:43,

tab. 2, fig. 5 [new synonymy].

This is a somewhat variable species in several respects and it is understandable that varieties have

been named. Specimens are always conspicuously

marked in black and white, the lighter individuals

being those referred to by Champion as the typical

nodulosus. Individuals with a greater proportion of the

dorsal surface black in color, Champion called variety

sallaei, with variety verrucosus somewhat intermediate

between the two.

The formation of the transverse callus at the apex

of the terminal abdominal sternum varies from simply

emarginate anteriorly to almost completely divided

into two distinct lobes. The antennal fossae are

appendiculate and the ventral surface of the pronotum

is usually smooth, but occasionally may have several

rather coarse tubercles.

Several unusual specimens are worthy of special

mention. In one from Santo Domingo, Oaxaca

(BMNH), there are numerous strong tubercles on the

ventral surface of the pronotum as in nervosus Solier;

the ventral surface is largely white and the femora are

white except at the extreme base and apex. Another

specimen exhibiting the same modifications is from 2

miles south of Tuxtla Gutierrez, Chiapas (UCal). A

specimen from Chilpancingo, Guerrero (MCZ), is

almost completely black and the dorsal surface is

smooth. This is probably an old abraded individual.

I have studied 16 of Champion's specimens, including both of the forms which he described, and am

convinced that only one species is involved. Champion

studied the type of Z. variolosus Sturm and pronounced it identical with this species.

MEASUREMENTS.—Length 14.1-28 mm; width

5.5-11.2 mm.

SPECIMENS EXAMINED.—137 from the localities that

follow. MEXICO: El Zapotal, 2 miles south of Tuxtla

Gutierrez in Chiapas; Acapulco, Chilpancingo, 38

miles south of Iguala, Omilteme and Taxco in Guerrero; San Miguel and Zacualtipan in Hidalgo; Ajijic

and Guadalajara in Jalisco; Mexico, Distrito Federal;

16 miles south of Arteaga in Michoacan; Cuernavaca

and Tejalpa in Morelos; Compostela and Ixtlan del

Rio in Nayarit; Calpulalpan, Juquila, Oaxaca, and

Santo Domingo in Oaxaca; 82 km northeast of Tehuacan in Puebla; 34.1 miles north and 40 miles south of

Tamazunchale in San Luis Potosi; 14 miles east of

Landa de Matamoros in Queretaro; Hermosillo in

Sonora; Canon de Santa Anna, Rancho La Mariposa,

Muzquiz in Coahuila; Tampico in Tamaulipas;

Jalapa, Orizaba, and Presidio in Veracruz. Specimens

were collected in every month except November at

altitudes of from 5,200 to 6,800 feet.

Zopherus nodulosus haldemani Horn,

new combination

PLATE 2: FIGURE 12

Zopherus haldemani Horn, 1870:271.—Champion, 1884:43.

—Casey, 1907b: 463.

This subspecies is apparently a continuation of the

trend from the dark {nodulosus) to the lighter

(sallaei) form. This is the lightest form of the series

and, as Casey (1907b: 463) remarks, is rather constant in coloration. The sutural rows of black blotches

are contiguous and form a continuous black band from

base to apex. There is also an irregular longitudinal

black band on the pronotum. The lateral areas of

both pronotum and elytra are predominately white.

Because the constant pattern of coloration appears

to be geographic, it seems advisable to retain the

trinomial for this and the previous taxon. Most of the

specimens seen from north of the Rio Grande River

are readily separable from those from Mexico.
NUMBE R 10 8

MEASUREMENTS.—Length 14.6-29.4 mm; width

5.7-11.3 mm.

SPECIMENS EXAMINED.—379 from the localities that

follow. UNITED STATES. TEXAS: Austin, Bastrop,

Benchley, Berclair, Brenham, Brownsville, Brownwood, Calvert, Camp Bullis, Camp Swift, College

Station, Columbus, Corpus Christi, Cypress Mill,

Deckers Prairie, Dime Box, Fort Sam Houston,

Galveston, Handley, Hearne, Kerrville, Lackland Air

Force Base, Lexington, Luling, Montell, New Braunfels, Palmetto State Park, Roma, Rock Island, Round

Mountain, Sabinal, Salado, San Antonio, Seagraves,

Taylor, 15 miles north of Temple, Twin Sisters,

Uvalde, Victoria, Waco, Dimmitt County, Eastland

County, Gillespie County, Grimes County, Harris

County, Limestone County, Montague County.

MEXICO. NUEVO LEON : Horsetail Falls, 25 miles south

of Monterrey; Chipinque Mesa, near Monterrey. SAN

LUIS POTOSI: El Salto Falls. DURANGO: 5 miles southwest of Cuencame.

I have seen specimens labeled "Guatemala," "California," "Monticello, Florida," and "Louisiana" and

am inclined to discount all except possibly the latter.

Specimens have been taken in every month except

February and December.

4. Zopherus jourdani Salle

PLATE 2: FIGURE 13

Zopherus jourdanii Salle, 1849:301, pi. 8: fig. 4.

Zopherus mexicanus (auct., nee. Gray).

Zopherus costaricensis Champion, 1884:40, tab. 2, fig. 1

[new synonymy].

This species may be separated from Z. jansoni by the

rounded and less prominent elytral base and in not

having the sides of the terminal elytral tubercles in

contact with the lateral margins; otherwise these two

species are quite similar in coloration and in having

distinctly bituberculate elytral apices. Both have a

pronounced dark band on the anterior pronotal

margin, which in jourdani is almost always crescent

shaped (rounded posterior margins) and in jansoni

is rectangular. This character is fairly reliable but not

without exception.

Coloration is variable. Specimens from Mexico have

larger black elytral tubercles than those from farther

south. The lightest specimens are from Costa Rica and

it is to this form that Champion assigned the name

costaricensis. A series of 24 specimens from 6 miles east

of San Cristobal, Chiapas, Mexico, collected 6 and 22

July 1957 by J. A. Chemsak and B. J. Rannells (CAS)

are the darkest I have seen and are quite uniform

througout the series. The disc of the pronotum is

almost entirely black as is the central portion of the

elytra.

In specimens from Costa Rica the black areas of

the elytra (tubercles and spots) are smaller and more

numerous, and the raised areas of the pronotum are

for the most part surrounded by white, giving the

entire dorsum a decided salt-and-pepper appearance.

In intermediate areas, specimens run the gamut

between the two extremes mentioned above. Specimens

from Guatemala and Honduras may indicate clinal

variations; however, two specimens from Chiapas,

Mexico, are identical in coloration to those typical of

Costa Rica. Considering the small number of specimens available from Honduras, Guatemala, and

Nicaragua, it seems best not to speculate upon the

meaning of this variability.

The ventral surface of the pronotum is very densely

and strongly tuberculate, and the antennal fossae,

while usually more less appendiculate apically, are

frequently rounded or truncate. Occasionally the

fossae will be appendiculate on one side and rounded

on the other side of a single individual.

Salle (1849) states that the natives of Guatemala

call this insect "Cameleon" in reference to its ability

to go for long periods of time without food. In Costa

Rica it is known locally as "Duerme Nino" because

of its deathfeigning behavior when disturbed.

MEASUREMENTS.—Length 17.7—36 mm; width

6.1-12 mm.

SPECIMENS EXAMINED.—187 from the localities that

follow. BRITISH HONDURAS: Punta Gorda, Rio

Sarstun. COSTA RICA: Alajuela, Azahar de Cartago,

Cachi, Coronado, Guadalupe, La Palma, Monte

Redondo, Monteverde, Navarro, Pacayas, Puntarenas,

Rio Sucio, Salinas, San Francisco, San Jose, Santa

Cruz, Volcan Irazu. EL SALVADOR: 10 km west of

Santa Ana, Santa Tecla, Sonsonate. GUATEMALA:

Capetillo, Chocoyos, Guatemala City, Panajachcl.

Totonicapan, Yepocapa. HONDURAS : El Zamorano,

Las Limas, Rosario, San Luis, Tegucigalpa, Yoro.

MEXICO: 8 km south of Bochil, Chenalho, 10 miles

northwest of Comitan, Pinca La Isle. Montidello Lake.

4.6 miles north of Ocozocuautla, 6 miles east of San

Cristobal, Tapachula, 3 miles southeast of La Trinitaria (all in Chiapas) ; Oaxaca in Oaxaca; Jalapa in
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Veracruz. NICARAGUA: 5 miles south of Jinotega,

Managua, Volcan Vie jo.

Specimens have been collected in every month at

altitudes ranging from 2,500 to 9,800 feet. It has been

taken on pine in Honduras and data accompanying

Costa Rica specimens are as follows: "under bark,"

"on balsa," "on tree trunk."

5. Zopherus jansoni Champion

Zopherus jansoni Champion, 1884:39, tab. 2, fig. 2.

This is a species of more slender body proportions

than jourdani. Differences between the two are summarized under the latter species.

MEASUREMENTS.—Length 19.7—27 mm; width

6.2-9.0 mm.

SPECIMENS EXAMINED.—22 from the localities that

follow. COSTA RICA: La Caja, Lindora Farm, Pacayas,

Pozo Azul de Pirris, Puntarenas, San Jose. NICARAGUA :

Chontales, 5 miles south of Jinotega. Specimens have

been taken in January, April, June, November, and

December.

6. Zopherus mexicanus Gray

PLATE 2: FIGURE 14

Zopherus mexicanus Gray, 1832:796, pi. 50: fig. 5; pi. 69:

figs. 4a-g.—Champion, 1892:489.

Zopherus maculatus Champion, 1884:41.

So few specimens of this species have been seen that

I can add nothing to what Champion has already

reported. Champion himself, after studying Gray's type

of mexicanus, placed his species maculatus in synonymy with it and I have accepted his decision.

Zopherus mexicanus looks very much like laevicollis

but has smaller, better defined apical tubercles of the

elytra, and the ventral callus of the terminal abdominal

sternum is feebly arcuate rather than trilobed (Figure

4). Specimens of the small series available vary considerably in coloration, from largely white to one

which is completely black (Linares in Nuevo Leon,

[CAS]).

MEASUREMENTS.—Length 14.6-24.2 mm; width

5.1-9.5 mm.

SPECIMENS EXAMINED.—9 from the localities that

follow. MEXICO: Cuernavaca in Morelos (CU);

Linares in Nuevo Leon (CAS); Zacualtipan in Hidalgo (BMHN); State of Jalisco (BMNH).

7. Zopherus angulicollis Champion

PLATE 3: FIGURE 15

Zopherus angulicollis Champion, 1884:42, pi. 2; fig. 6.

This species, as indicated by the key, is similar to

mexicanus, xestus, and laevicollis in having large,

flattened tubercles on the elytral apex. In angulicollis

these tubercles are better developed than in the other

three species and are almost contiguous at their bases

(Figure 2). The prominent sutural ridge that separates

the elytral tubercles in the other three species is not at

all developed in angulicollis. The development of the

ventral callus of the apical abdominal sternum is intermediate between the simply arcuate form of mexicanus

and the strongly trilobed condition of laevicollis and

xestus. It is not at all like guttulatus ( = concolor) as

suggested by Champion.

MEASUREMENTS.—Length 17.0—18.7 mm; width

5.9-7.1 mm.

SPECIMENS EXAMINED.—I have studied the unique

type from Pinos Altos, Chihuahua, Mexico (BMNH),

and can report one additional specimen labeled as

follows: Tres Rios, Chihuahua, Mexico, 29 June 1958,

Tanner-Robinson (in collection of Dr. Vasco M.

Tanner, Brigham Young University).

8. Zopherus laevicollis Solier

PLATE 3: FIGURE 16

Zopherus laevicollis Solier, 1841:46.—Champion, 1884:40.

Zopherus laevicollis var. venosus Champion, 1884:40;

1892 :488. [new synonymy].

Zopherus tuberculatus Champion, 1884:41, tab. 2, fig. 3.

[new synonymy].

Zopherinus limbatus Casey, 1907a:37 [new synonymy].

This is an extremely variable species in dorsal coloration, luster, and sculpture. Coloration ranges from

completely black individuals to specimens in which

the dorsal surface is in greater part white, with

prominent black, flattened tubercles. In some specimens the tubercles are large and numerous and are

arranged in definite longitudinal rows. In others the

tubercles tend to coalesce and form wavy transverse

plicae. A few individuals have the tubercles well

separated and surrounded by the white ground color.

Two specimens are entirely black, without trace of

white markings whatsoever. Surface luster ranges

from dull to distinctly shining. This species may be
NUMBE R 10 8 11

distinguished from all other species of Zopherus except

xestus by the peculiar formation of the ventral callosity of the terminal abdominal sternum. In both

laevicollis and xestus there is a prominent ridge projecting cephalad from the normal bilobed callosity,

causing this structure to appear trilobed as described by

Solier.

After studying a good series of Champion's specimens of venosus and specimens which he determined

as laevicollis, I concur with his earlier (1884) conviction that they are not separable. I have seen only

one of Champion's two specimens of tuberculatus from

Santo Domingo, Oaxaca, Mexico, mentioned in the

Biologia and see no reason for separating that species

from laevicollis.

MEASUREMENTS.—Length 13.7—22.4 mm; width

4.9-8.2 mm.

TYPES.—Zopherinus limbatus Casey. The Casey

collection contains 6 specimens bearing the USNM

type number 46366. All are from Amecameca,

Morelos, Mexico, collected by Hoge. I have selected

and labeled the first one of the series a lectotype. All

of the Casey specimens fall well within the range of

variation observed in the numerous examples of this

species that were studied. Casey had one specimen

determined as laevicollis, also from Amecameca. To

me it appears identical to those he called limbatus.

One of his specimens of limbatus bears a "Z. venosus"

label. Apparently Casey saw so few specimens of

laevicollis that he had no concept of its range of

variability.

SPECIMENS EXAMINED.—216 from the localities that

follow. MEXICO. CHIAPAS: 10 miles north of Tuxtla

Gutierrez. CHIHUAHUA: 30 miles west of Belleza.

COAHUILA: 33 miles southeast of Saltillo. DISTRITO

FEDERAL: 15 miles south of El Guarda, Mexico City,

Tacubaya. DURANGO: 6 miles northeast of El Salto,

San Miguel, Tepehuanes. GUERRERO: Acapulco,

Mochitlan. HIDALGO: Zacualtipan. JALISCO: Mazamitla. MEXICO: Amecameca, Chapingo, Otumba.

MICHOACAN : Uruapan. MORELOS : Cuernavaca.

NUEVO LEON: Galeana, 10 miles east of San Roberto.

OAXACA: Juquila Mixes, La Parada, Ruinas de Mitla,

Salina Cruz, Santo Domingo, 17.2 miles northwest of

Tehuantepec, Km 180-212 Puerto Escondido Highway, 22.3 miles west northwest of Zanatepec. PUEBLA:

3 miles northeast of Zacatepec. TLAXCALA: Apizaco.

VERACRUZ : Jalapa, Las Vigas, Tlapacoyan. Champion

listed it from the following localities: Sierra de San

Miguelito (San Luis Potosi), San Andres Chalchicomula (Puebla), Juquila and Calpulalpan (Oaxaca ). Specimens have been taken in every month

except March.

9. Zopherus xestus, new species

PLATE 3: FIGURE 17

DESCRIPTION.—Holotype, sex undetermined. Elongate, moderately robust, dull, velvety black in color.

Head entirely black, epistomal margin evenly and

deeply emarginate, dorsal surface minutely and very

sparsely punctate, punctures not evidently setigerous;

ventral surface smooth except for a few feeble punctures on anterolateral margins; mentum transversely

trapezoidal, apical margin bisinuate with prominent

anterior angles, finely and very sparsely punctate.

Pronotum in dorsal view exactly as broad as long,

sides strongly and evenly arcuate, with a slight prominence above apex of antennal fossa and five rather

prominent tubercles on basal half of lateral margins;

sinuate just before base, apical margin broadly and

shallowly emarginate, the angles bluntly rounded,

basal margin slightly rounded, thickened medially,

angles rather sharply rectilinear; surface black except

for feeble indications of white or gray lines parallel to

and just within lateral borders, very minutely and

sparsely punctulate; ventral surface densely opaque

with a few small, widely scattered punctures on prosternum; antennal fossae rounded apically. Elytra

slightly less than twice as long as broad, sides evenly

rounded, base broadly and deeply emarginate, as wide

as pronotal base; humeri acute and prominent, basal

margin distinctly thickened and reflexed; apical

tubercles smooth, shiny, separated from lateral margin

by dull ground color and a few small punctures forming a fine groove; sharply separated from raised sutural

margin by deep grooves; surface practically smooth,

with a series of shallow, irregular, and poorly defined

lines producing a wavy pattern over entire surface;

very minutely and sparsely punctulate; black in color,

with feeble indications of white or gray lines forming

a fine network in depressions along lateral border.

Mesosternum and metasternum densely opaque,

minutely and sparsely punctulate; abdominal sterna

likewise sparsely punctate, except on caudal margins

of segments 3 and 4 where punctures are more numerous and closely spaced; ventral callus of terminal

abdominal sternum trilobed, with the median lobe
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rounded and prominent, the usual patch of yellowish

hair present. Legs coarsely but sparsely punctured.

VARIATION.—The specimen selected as the holotype

is almost intermediate in intensity of elytral sculpturing. Some specimens are smoother and show a lesser

amount of white on lateral portions of the pronotum

and elytra. This I take to be a result of aging and

wear. This species is most like laevicollis Solier and is

the only one north of the Rio Grande River which has

the trilobed ventral abdominal callus and the large

flattened tubercles at the apex of the elytra.

MEASUREMENTS.—Holotype: Length 22.4 mm;

width 8.2 mm. Paratypes: Length 16.8—24.8 mm;

width 6.0-8.8 mm.

SPECIMENS EXAMINED.—Holotype: Brewster County, Texas, Big Bend National Park, The Basin of

Chisos Mountains, 8 August 1962, C. A. Triplehorn

(OSU). 37 paratypes from same locality, 5 May—3

September, collected by C. A., W. E., B. A., and B. W.

Triplehorn, D. J. and J. N. Knull, Howden and

Becker, W. R. M. Mason, R. and A. Graves, J. A.

Brubaker, R. L. Berry and F. J. Moore, R. and L.

Hamilton and Howard V. Weems, Jr., (OSU, CNC,

FSCA, RLB, TAM, UCR) ; (1) Big Bend National

Park, Green Gulch (5700'), 24 July 1968, J. E. Hafernik (TAM); (1) Big Bend National Park, Panther

Junction (4,000 feet) 29-30 April 1959, Howden and

Becker (CNC); (1) Davis Mountains, Jeff Davis

County, Texas, 19 June 1958, D. J. and J. N. Knull

(OSU).

10. Zopherus solieri, new species

Zopherus elegans Champion (nee Horn), 1892:489.

DESCRIPTION.—Holotype, sex undetermined.

Moderate in size, robust, black and white, dull in

luster. Head entirely black, epistomal margin evenly

and rather deeply emarginate, frons somewhat swollen

in front of eyes, dorsal surface finely and sparsely

punctate, each puncture with a short, inconspicuous,

yellowish seta; ventral surface smooth except for a

few coarse punctures; mentum broadly transverse,

apical margin bisinuate with angles lobed and prominent, coarsely and sparsely punctured. Pronotum in

dorsal view slightly broader than long, sides strongly

arcuate, somewhat angulate above apex of antennal

fossae, briefly sinuate just before base, apical margin

broadly and shallowly emarginate, the angles bluntly

rounded, basal margin feebly rounded, uniformly

thickened, angles obtusely rounded; surface, except for

lateral borders, black and alutaceous with moderately

coarse, shallow, widely spaced punctures; white lateral

borders occupying about one ninth the pronotal width

on each side, inner border poorly defined; numerous,

dense, shining black tubercles prominent within white

borders; ventral surface black; antennal fossae rounded apically with angle somewhat upturned; prosternum

coarsely and densely punctured, prosternal process

shallowly excavate between coxae, ventral surface of

prothorax with a few scattered, coarse punctures. Elytra almost twice as long as broad, sides evenly rounded,

base broadly and shallowly emarginate, slightly wider

than pronotal base, coarsely tuberculate with humeri

prominent; apical ridges black, moderately swollen,

oblique, very narrowly in contact with raised suture

where it passes between them, mostly distinct from lateral margin except at extreme apex; surface strongly

sculptured with irregular flattened tubercles which are

confusedly arranged and tend to coalesce, forming

wavy transverse plicae, punctures extremely fine and

sparsely distributed; median seven ninths black in color

and alutaceous, outer borders white, densely spotted

with black, fairly regular, raised tubercles which frequently are contiguous, inflexed portion entirely black,

finely and sparsely punctured. Mesosternum and metasternum coarsely and densely punctured, abdominal

sterna sparsely punctured, with punctures of varying

size, ventral callus of terminal abdominal sternum

with an acutely rounded and prominent median lobe

produced cephalad, the usual patch of yellowish hairs

present. Legs entirely black except for the dense rows

of yellowish hairs, coarsely and conspicuously punctured.

VARIATION.—The greatest variation observed is

in the extent of the white borders of the elytra. The

white in some of the specimens occupies as much as

one quarter of the width of the elytra on each side,

with a few small white patches on the disc. Not all

of the specimens have the apex of the antennal fossae upturned; this character is subject to variation in

this as in several other species of Zopherus.

MEASUREMENTS.—Holotype: Length 18.2 mm;

width 6.6 mm. Paratypes: Length 17.1—20.3 mm;

width 6.3-7.2 mm.

SPECIMENS EXAMINED. — Holotype and five paratypes, all from Villa Lerdo, Durango, Mexico, collected by Hoge (BMNH); one paratype, same data
NUMBE R 10 8 13

(ANSP, Horn collection 7216) ; one paratype, same

data (MCZ), obviously obtained through exchange

from Champion.

11. Zopherus championi, new species

PLATE 3: FIGURE 18

Zopherus elegans Champion (nee Horn), 1892:489, tab. 22,

fig. 7.

DESCRIPTION.—Holotype, sex undetermined. Short,

stout, black and white, dull in luster. Head entirely

black; epistomal margin evenly and feebly emarginate, dorsal surface coarsely but shallowly and sparsely

punctured, each puncture with a short, coarse, yellowish seta, ventral surface smooth except for a few widely spaced punctures laterally; mentum broadly transverse, apical margin bisinuate with angles lobed and

prominent, coarsely and sparsely punctured. Pronotum

in dorsal view just slightly broader than long, sides

strongly arcuate, somewhat angulate above apex of

antennal fossae, abruptly and briefly sinuate just

before base, apical margin broadly and shallowly

emarginate, the angles bluntly rounded, basal margin

slightly rounded, thickened medially, angles obtusely

rounded; surface, except for lateral borders, black

and alutaceous with widely spaced, fairly coarse but

extremely shallow punctures, white lateral borders

occupying less than one seventh the pronotal width

on each side, inner border rather irregular, a few

very small black spots within white margins, these

more numerous anteriorly; ventral surface black; antennal fossae rounded apical ly, prosternum coarsely, deeply, and densely punctured, prosternal process

shallowly excavate between coxae, ventral surface of

prothorax entirely smooth, impunctate. Elytra about

two thirds as wide as long, sides evenly rounded, base

broadly and deeply emarginate, slightly wider than

pronotal base, humeri acute and prominent; apical

ridges black, moderately swollen, oblique, broadly in

contact with raised suture where it passes between

them, also continuous with apical margin; surface

feebly irregular without distinct tubercles, punctures

coarse, shallow, and very sparsely distributed, median

three fifths black in color and alutaceous, outer borders white, irregularly spotted with black blotches, inflexed portion entirely black, coarsely and sparsely

punctured. Mesosternum and metasternum coarsely

and irregularly punctured; abdominal sterna coarsely and densely punctured medially but practically

impunctate laterally; ventral callus of terminal abdominal sternum trilobed with the median lobe acutely

rounded and prominent, the usual patch of yellowish

hair present. Legs entirely black, except for the usual

pattern of yellowish hairs, conspicuously punctured.

VARIATION. — The elytral sculpturing in some of

the specimens is somewhat more irregular than in the

type, with an approach to distinctly flattened tubercles.

The series is, for the most part, remarkably constant

in coloration and general appearance.

MEASUREMENTS.—Holotype: Length 13.3 mm;

width 5.3 mm. Paratypes: Length 11.9—15.1 mm;

width 4.3—5.6 mm.

SPECIMENS EXAMINED. — Holotype and four paratypes from Villa Lerdo, Durango, Mexico, collected

by Hoge (BMNH). The holotype is the specimen

from which Champion's illustration was prepared.

Paratypes: (1) Huasteca Canyon, near Monterrey,

Nuevo Leon, Mexico, 11 July 1963, H. F. Howden

(CNC) ; (1) same locality and date, R. H. Arnett,

Jr., and E. R. Van Tassell (FSCA) ; (1) Sanderson,

Texas, 7 August 1959, R. B. Selander and J. C. Schaffner (JMC). One additional specimen, not designated

a paratype because the white pattern cannot be seen:

Sanderson, Texas, 6 June 1956, H. and A. Howden

(OSU).

12. Zopherus tristis LeConte

PLATE 3: FIGURE 19

Zopherus tristis LeConte, 1851:130.—Horn, 1867:161;

1870:272;1894:347.

Zopherodes tristis, Casey, 1907a: 38.—Blaisdell, 1923:243;

1931:113.

Zopherodes aequalis Casey, 1907a:38 [new synonymy].

Zopherodes variabilis Casey, 1907b:464 [new synonymy].

Zopherodes incrustans Casey, 1907b:464 [new synonymy].

This is one of the easiest species of Zopherus to recognize, but one of the most difficult to characterize. The

body is subcylindrical, greatly thickened dorsoventrally, with the elytral margins parallel throughout a

greater portion of their length and abruptly constricted at the base and apex. The apical declivity is more

abrupt and steeper than in any other species. The

elytral tubercles are usually rather flattened with their

hind margins somewhat truncate. Most specimens

have the pronotum coarsely punctured, but several

have coarsely granulate punctures as in granicollis;

the prosternum is coarsely punctured. Sometimes the
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

flanks of the pronotum and elytra are encrusted with

a whitish substance, and this may frequently be restored by soaking the specimen in ether. It appears

to be of no taxonomic significance.

Casey (190 7b: 467) considered regarding the species he named aequalis, incrustans, and variabilis as

subspecies of tristis. I have examined the types and

they all fall well within the range of variation exhibited by tristis.

MEASUREMENTS. — Length 10.5-22 mm; width

3.9-7.6 mm.

TYPES. — Zopherus tristis LeConte (not seen).

Type locality: "in deserta fluminis Colorado." Zopherodes aequalis Casey. Holotype (USNM 46350).

Arizona (Gila Valley), Dunn. Zopherodes variabilis

Casey. I have selected and so labeled the first of six

specimens under this name in the Casey collection as

lectotype. All were collected in the Baboquivari

Mountains, Arizona, by F. H. Snow and bear a

USNM type number 46351. This series is quite variable in size and sculpture, but one of them (the third)

is a perfect match for the type specimen of aequalis

Casey. Zopherodes incrustans Casey. Holotype

(USNM 46349). Tucson, Arizona, F. H. Snow.

SPECIMENS EXAMINED. — 276 from the localities

that follow. UNITED STATES. ARIZONA: Ajo, Baboquivari Mountains, Buckeye, Bumble Bee, Chandler,

Florence, Granite Reef, Mesa, Nogales, Organ Pipe

Cactus National Monument, Parker, Phoenix, Prescott, Riverside, Safford, San Carlos Lake, Santa

Catalina Mountains, Tacna, Tempe, Tucson, Vail,

Wellton, 12 miles east of Wenden, Wickenburg,

Yuma. CALIFORNIA: Blythe, Borrego, Calexico, Caliente, Coachella Valley, Carrizo Springs, Holtville, Imperial, Indio, Laguna, Palm Springs, Palo Verde,

Thermal, Winterhaven, Yuma. TEXAS : One specimen

with State label only, probably in error. MEXICO.

BAJA CALIFORNIA: Comondu, Loreto, Mulege, 1 mile

south of Puerto Escondido, 10 miles south of Punta

Prieta, San Domingo, 5 miles south of San Miguel,

25 miles south of Santa Rosalia. SONORA: Guaymas,

Hermosillo, Pesqueria. Specimens have been collected

in even- month except December.

13. Zopherus concolor LeConte

PLATE 3: FIGURE 20

Zopherus concolor LeConte, 1851:130.

Zopherus guttulatus Horn, 1867: 160 [new synonymy].

Zopherodes concolor, Casey, 1907a: 38.

Zopherodes guttulatus, Casey, 1907a: 39.

Zopherodes morosus Casey, 1907b:465 [new synonymy].

This species is characterized by its coarse elytral

sculpturing, consisting of large, rounded tubercles

which are usually contiguous at their bases and frequently form transverse wavy plicae. The pronotal

punctures are sparse and simple, at least never forming

rough granules, and the prosternum is usually roughly sculptured with punctures rather than raised tubercles. The form described by Horn as guttulatus has a

somewhat linear arrangement of the elytral tubercles,

but there are many intermediates connecting this form

with the more typical sculpture. Casey's specimens of

morosus, including the lectotype, are very typical concolor.

In several specimens it has been possible to bring

out a white lateral border, almost identical to that of

elegans, by prolonged soaking in ether. Specimens of

elegans may be distinguished by the much finer elytral

sculpture, the coarsely granulate pronotal punctures,

and the strongly tuberculate prosternum.

MEASUREMENTS. — Length 12.4-22.8 mm; width

4.5-8.2 mm.

TYPES. — Zopherus concolor LeConte. Holotype

(not seen). Near Santa Fe, New Mexico.

Zopherus guttulatus Horn. Lectotype (ANSP 3921).

Texas (southwestern) ; one paratype labeled same as

lectotype.

Zopherodes morosus Casey. Three specimens from

Cloudcroft, New Mexico, 9,000 feet, W. Knaus, represent this species in the Casey collection. The first specimen has been selected and so labeled a lectotype by

me. Both specimens also bear a USNM type number

46352.

SPECIMENS EXAMINED.— 121 from the localities

that follow. UNITED STATES. NEW MEXICO: Albuquerque, Bandelier National Monument, Bent, Cloudcroft, Conchas Dam, Coolidge, Corona, Duran, High

Rolls, Hot Springs, Jemez Springs, Las Vegas, Mayhill, Mescalera Reservation, Omega, Romeroville,

Ruidoso, Santa Fe, Tecolote. TEXAS : Davis Mountains, Lake Buchanan (Llano County), Pine Springs

(Culberson County), 20 miles northwest of San

Antonio, 16 miles south of Sheffield (Terrell County),

35 miles west of Toyah (Culberson County), Real

County. One specimen from Nogales, Arizona (WR),

but data may not be authentic. Specimens have been
NUMBE R 10 8 15

collected from 11 June to 19 December, but by far

the largest numbers were taken in June.

14. Zopherus gracilis Horn

PLATE 3: FIGURE 21

Zopherus gracilis Horn, 1867:162; 1870:272.

Zopherodes pudens Casey, 1907a:40 [new synonymy].

Zopherodes caudalis Casey, 1907a:41 [new synonymy].

Zopherodes lugubris Casey, 1907a: 41; 1911:254 [new

synonymy].

Zopherodes pruddeni Casey, 1907a:41 [new synonymy].

Zopherodes luctuosus Casey, 1907a:41 [new synonymy].

Zopherodes elongatus Casey, 1907b:466 [new synonymy].

Zopherodes geminatus Casey, 1907b:467 [new synonymy].

This is the smoothest known member of the genus

Zopherus. Typically, the elytra are smooth but may

have fine, remotely spaced punctures and scattered,

randomly arranged wavy incised lines. This species

occurs from the Grand Canyon area of Arizona to

southeastern Arizona, southwestern New Mexico, and

into at least northern Sonora, Mexico. It is especially

abundant in the Chiricahua Mountain area, where

sizable series have been taken at night on bark of

pine.

Z. gracilis varies clinally from south to north in

the intensity of elytral sculpture and punctation. Specimens from southeastern Arizona (Chiricahuas) and

adjacent regions of New Mexico are almost invariably

perfectly smooth, whereas those from Flagstaff and the

south rim of Grand Canyon sometimes approach

uteanus in sculpture. It is thus not too surprising that

Colonel Casey proposed seven names for some of the

variations observable in this species. It is rather unforgivable that he did not recognize Horn's gracilis.

Four of Casey's "species" are represented only by the

unique types, two others by only two specimens each,

and the remaining one by three specimens. It is obvious that he was describing specimens rather than

species.

The apical ridges of the elytra are poorly denned

and only slightly swollen. They are in more or less

continuous contact with the lateral and apical margins of the elytra and are separated from the apical

sutural elongation by a deep groove except at the

extreme apex. The ventral callus of the terminal abdominal segment is broadly arcuate, with an acutely

rounded and prominent median lobe directed cephalad. The prosternum is coarsely tuberculate and

the antennal fossae are rounded apically.

MEASUREMENTS. — Length 11.6—21.7 mm; width

3.8-8.4 mm.

TYPES. — Zopherus gracilis Horn. Arizona (in

the neighborhood of Fort Whipple, fide Horn). I

have not seen the type which is stated by Horn to be

in the LeConte collection; however, I have seen four

specimens from Horn's collection.

Zopherodes pudens Casey. Holotype (USNM

46346) Ari[zona]. This unique specimen represents

the maximum elytral sculpture in the Casey collection. It is rather typical of those found in the Flagstaff area.

Zopherodes caudalis Casey. Holotype (USNM

46363), Ari[zona]. A unique. The long terminal

grooves of the elytra are not really so striking as

Casey suggests.

Zopherodes lugubris Casey. This species is represented in the Casey collection by two specimens: the

first labeled "Oslar, Tucson, Arizona," designated as

lugubris in Casey's handwriting, and the second

labeled simply "Ariz." A note in the box, presumably

by L. L. Buchanan, states that "Casey evidently synonymyzed lugubris with pruddeni and then selected

two Arizona examples to represent the former." These

two specimens are so unlike in elytral sculpture that

I am surprised that Casey did not describe them as

distinct. Both, however, are typical gracilis. The type

locality was originally designated as "Grand Canyon

of the Colorado" but later changed by Casey (1911:

254) to "near Tucson." The original type of lugubris,

a unique, now stands under Z. pruddeni (see remarks

below).

Zopherodes pruddeni Casey. Three specimens stand

under this name in the Casey collection. I have selected and so labeled the first one in the series, said by

Casey to be from the Grand Canyon of the Colorado,

as lectotype. This specimen also has a USNM type

number 46361. The original description indicates that

this species was described from a unique specimen.

A second specimen under this name in Casey's collection and labeled the same way is doubtless the

original type of lugubris (USNM 463.62). A third

specimen, labeled simply "Ariz," is apparently not

from the same locality as the other two. All three are

alike in sculpture and exhibit no departure from

normal variation in this species.

Zopherodes luctuosus Casey. Holotype (USNM

46365), Ari[zona]. A unique which comes closest to
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Horn's concept of gracilis among all of the specimens

in Casey's collection.

Zopherodes elongatus Casey. Casey had two specimens under this name, both labeled Ari[zona] and

USNM type number 46364. I hereby designate, and

have so labeled the first of these, a lectotype. This

form is no more slender than others I have seen.

Zopherodes geminatus Casey. Holotype (USNM

46355), Ari[zona], Fort Apache, according to Casey.

The twinning of elytral punctures upon which Casey

largely bases this species is not a unique character

for I have seen it in occasional individuals from various parts of Arizona and New Mexico.

We may arrange the forms described as species by

Casey (and here reduced to synonyms of gracilis) in

order of increasing intensity of elytral sculpture as

follows: luctuosus (practically smooth), geminatus,

lugubris, caudalis, pruddeni, elongatus, and pudens

(actually somewhat tuberculate). As stated before,

these form a fairly constant cline from south to north.

SPECIMENS EXAMINED. — 591 from the localities

that follow. UNITED STATES. ARIZONA: Ash Fork,

Bright Angel Camp, Cameron, Chiricahua Mountains,

Crown King, Flagstaff, Fort Grant, Fort Valley, Galiuro Mountains, Globe, Graham Mountains, South

rim of Grand Canyon, Huachuca Mountains, Jerome,

Mesa, Morrison, Nogales, Oak Creek Canyon, Oracle,

Patagonia, Peach Springs, Pine, Portal, Prescott,

Riverside, Ruby, Santa Catalina Mountains, Santa

Rita Mountains, Seligman, Seven Springs, Sierra

Ancha Mountains, Tombstone, Williams. NEW MEXICO: Lordsburg, Rodeo, Silver City. MEXICO. 10

miles east of Cananca in Sonora. Specimens have

been collected from April to December.

15. Zopherus uteanus (Casey), new combination

PLATE 4: FIGURE 22

Zopherodes uteanus Casey, 1907a: 40.

Zopherodes mormon Casey, 1907a:40 [new synonymy].

This species is very similar in appearance to Z.

opacus Horn in its uniformly dull dorsal surface luster.

Some specimens also approach Z. granicollis ventriosus in elytral sculpture but may be distinguished

by the fine, sparse punctation of the head. In granicollis ventriosus the punctures of the head are very

coarse and dense. The elytral sculpturing consists

of small, shiny tubercles separated by dull interspaces.

Typically these tubercles are so densely spaced that

their bases are more or less contiguous, but sometimes they are smaller and more distantly spaced, approaching the situation in opacus. It is this latter

variation that Casey named mormon. Enough intermediate forms have been studied to convince me that

only one species is involved. In all specimens examined

the pronotum is coarsely and densely granulate-punctate.

MEASUREMENTS. — Length 14.0-22.0 mm; width

4.6-8.0 mm.

TYPES.—Zopherodes uteanus Casey. Holotype

(USNM 46344), southwestern Utah.

Zopherodes mormon Casey. Holotype (USNM

46345), Utah (no specific locality).

Each of the above is represented in the Casey collection by only the unique holotype, and this is

another instance in which Casey described specimens

and not species.

SPECIMENS EXAMINED. — 81 from the localities that

follow. UNITED STATES. ARIZONA: Grand Canyon

National Park (North Rim), Kaibab National Forest.

CALIFORNIA: Granite Mountains near Cottonwood

Springs (San Bernardino County). NEVADA: Baker,

Ely, Kyle Canyon in Charleston Mountains (Clark

County), Lee Canyon in Spring Mountains (Clark

County). UTAH: Arches National Monument, Bears

Ears (Elk Ridge), Beaver, Bryce Canyon, Dividend,

Eureka, LaSal, Leeds, Lehi, Marysvale, Montezuma

Creek (San Juan County), Natural Bridges National

Monument, Nephi, Paria, Parowan, Pine Valley,

Saint George, Stockton, Zion National Park. Collection

dates are from 15 April to 11 September. I have collected this species under bark of dead pines in Arches

National Monument and under chips of cut pine on

the ground in Bryce Canyon.

16. Zopherus granicollis granicollis Horn

PLATE 4: FIGURE 23

Zopherus granicollis Horn, 1885 :160.

Zopherodes induratus Casey, 1907a:39 [new synonymy].

Zopherodes californicus Casey, 1907b: 466 [new synonymy].

Zopherodes prominens Casey, 1924:305 [new synonymy].

The elytra of this robust species are typically coarsely

tuberculate, the tubercles closely spaced and more or

less contiguous at their bases. The pronotum is coarsely

and densely, frequently granulately punctured, and
NUMBER 10 8 17

the punctures of the head are coarse, deep, and closely

spaced. The dorsal surface is dull in luster.

This, the nominate subspecies, inhabits the mountains of southern California, and northern Baja California. I have seen only one specimen from Arizona

and one from Nevada (Casey's type of Z. induratus

from Las Vegas). An interesting and rather uniform

series of 14 specimens from Death Valley National

Monument was studied. These appear to be somewhat intermediate between this and the following

subspecies (ventriosus), the elytral tubercles being a

little smaller and more widely spaced. The lateral

elytral tubercles, however, are not transversely elongated as in typical ventriosus. I have seen this same

condition in series in which most individuals were

coarsely tuberculate and had assumed that it represented older specimens in which the elytra were

eroded. This cannot be the case with the Death Valley

specimens since they were reared from pine roots

(Pinus monophylla) by W. H. Tyson and pinned

immediately. More finely sculptured specimens of

this species may be separated from Z. uteanus by the

more robust body proportions and the coarsely and

densely punctured clypeus.

MEASUREMENTS. — Length 12.6-20.5 mm; width

4.4—7 .4 mm.

TYPES. — Zopherus granicollis Horn. Holotype

(ANSP 3922), California; a square yellow label indicating "northern part of Lower California."

Zopherodes induratus Casey. Holotype (USNM

46359), Julian, California. The unique type is a

typical specimen of granicollis; the only noteworthy

feature is the lack of a cephalad-projecting spur on

the ventral abdominal callus. This is unusual but

not unprecedented since I have seen several other

specimens with the same modification.

Zopherodes californicus Casey. Holotype (USNM

46357), Cal [ifornia]. In the unique type specimen,

the elytral tubercles are eroded and form transverse,

wavy plicae, a variation not uncommon in Z. granicollis.

Zopherodes prominens Casey. Holotype (USNM

46360), Las Vegas, Nevada, 26 May 1905, Tom

Spaulding. This specimen, also a unique, falls well

within the range of variation in this subspecies, the

pronotal punctures having perhaps a sparser arrangement than usual.

SPECIMENS EXAMINED.—126 from the localities

that follow. UNITED STATES. ARIZONA: Yuma

County. CALIFORNIA: Claremont, Cleghorn Canyon,

Crestline, Crystal Lake, Cuyamaca Rancho State Park,

Death Valley National Monument, Fillmore, Herron

Hill, Hesperia, Idyllwild, Joshua Tree National Monument, Julian, Lake Arrowhead, Littlerock, Mount

Wilson, Newhall, Palm Springs, Pasadena, Potrero,

Providence Mountains, San Bernardino Mountains,

San Gabriel Mountains, San Jacinto Mountains,

Santa Barbara, Santa Rosa Mountains. NEVADA: Las

Vegas. MEXICO. BAJA CALIFORNIA: Ensenada, Rumorosa, 2 miles south of El Toro (Sierra Juarez).

Specimens have been collected in every month except November and December.

Zopherus granicollis ventriosus (Casey),

new combination

PLATE 4: FIGURE 24

Zopherodes ventriosus Casey, 1907a: 39.

Zopherodes parvicollis Casey, 1907b: 466 [new synonymy].

This subspecies differs from the nominate subspecies

in having much finer elytral tubercles, with the lateral

ones always transversely elongate. The pronotal punctures are variable as in typical granicollis, varying

from coarsely muricate to granulate. Except for the

previously mentioned Death Valley specimens, I have

seen no intermediate specimens between the two subspecies, granicollis and ventriosus, apparently occupying reciprocal ranges. No specimens are available

from the broad area which separates the two.

MEASUREMENTS.—Length 14.5—21.0 mm; width

6.0-7.0 mm.

TYPES. — Zopherodes ventriosus Casey. Holotype

(USNM 46356), California (southern, fide Casey).

Zopherodes parvicollis Casey. Holotype (USNM

46358), Round Meadow Giant Forest, California,

Hopping.

Both of these are very similar in sculpture, the

principal difference being that the latter is very dull

in surface luster. Both are represented in the Casey

collection by uniques.

SPECIMENS EXAMINED. — 48 from the localities that

follow. UNITED STATES. CALIFORNIA: Cedar

Grove, Dalton Creek at 4,800 feet. Huckleberry Meadow, Kings River, Millwood, and Woods Creek in

Fresno County; Saline Valley in Inyo County; Kern
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

River Camp 3, Sequoia National Forest, and Mill

Potrero in Kern County; Bass Lake and Sugar Pine

in Madera County; Miami Ranger Station at 4—5,000

feet in Mariposa County; Colony Mill, Davenport,

Atwells Mill, Kaweah, Lloyd Meadow, Round Meadow Giant Forest, and Wells Mill in Tulare County.

Specimens have been collected from 3 April to 16

October.

17. Zopherus opacus Horn

PLATE 4: FIGURE 25

Zopherus opacus Horn, 1867:161; 1870:272.

Zopherodes opacus, Casey, 1907a: 40.

The elytral tubercles in this species are extremely

small and widely separated. The broad expanses

between tubercles are densely opaque and dull in

luster. The pronotal punctures are coarsely muricate

and closely spaced, and the prosternum is coarsely

and densely, sometimes rugosely, punctured.

MEASUREMENTS. — Length 13.0—20.3 mm; width

4.9-7.2 mm.

TYPES. — Holotype (ANSP 3923) Nevada (no

specific locality).

VARIATION. — In one of two specimens from Mono

County, California, the elytral tubercles are transversely elongate, with thin linear crests and a bit

closer together than usual. Horn had a similar specimen from Nevada, so I assume that this can be regarded as normal intraspecific variation. It should

be noted that Casey's concept of this species was

correct, although he had only one specimen in his

collection.

SPECIMENS EXAMINED. — 21 from the localities that

follow. UNITED STATES. CALIFORNIA: 9 miles west

of Lone Pine, Westgard Pass and Whitney Portal in

Inyo County; Convict Lake and Long Valley in Mono

County and Sage Flats (not located). NEVADA: Ruby

Substation, 10 miles southwest of Wells (6,000 feet)

in Elko County; Cody Basin and Reno in Washoe

County. UTAH : State label only. Specimens have been

taken from 25 February to 12 August.

Zopherus opacus is most like Z. uteanus Casey but

also bears some resemblance to Z. granicollis ventriosus. It displays by far the dullest surface luster

of any known member of the genus and should be

easily recognized.

18. Zopherus elegans Horn

PLATE 4: FIGURE 26

Zopherus elegans Horn, 1870:272.

Zopherodes otiosus Casey, 1907a:39 [new synonymy].

Zopherodes verrucipennis Casey, 1907b:465; 1911:254.

Zopherodes elegans, Casey, 1911: 254.

Zopherodes circumductus Casey, 1924:304 [new synonymy].

Zopherodes woodgatei Casey, 1924:305 [new synonymy].

Zopherodes otiosus var. verrucipennis, Casey, 1924:305.

This attractive and interesting species is easily identified by the whitish lateral borders of the pronotum

and elytra. The pronotum is always coarsely granulatepunctate, and the prosternum is distinctly tuberculate

rather than punctate. The elytra are sculptured

with rounded tubercles which vary somewhat in

size, shape, and arrangement. In some specimens

they are arranged in longitudinal rows (Casey's

woodgatei) and in some they are randomly scattered

(Casey's circumductus). Usually the tubercles are so

large that at least some of their bases are contiguous,

often forming transverse, wavy plicae. Occasionally

they are smaller and completely isolated from one

another, in which case there are usually much smaller

tubercles scattered within the more or less opaque

interspaces. The white areas surround the lateral

tubercles of both pronotum and elytra, causing the

black crests of the tubercles to stand out in bold relief. In 12 specimens it was impossible to produce the

white lateral areas, even by long periods of soaking

in several solvents. Except for the uniform black

color, these specimens are identical in sculpture and

body proportions to typical elegans. It is to this melanistic phase that Casey assigned the name Z. otiosus. I

see no reason to retain it as even a varietal name.

The 12 completely black specimens studied are from

the following localities: ARIZONA: Eagar, Pinal Mountains, White Mountains, 8-15 miles northeast of

Whiteriver. NEW MEXICO: Apache National Forest,

Fort Wingate ('ectotype and paratype of otiosus and

holotype of verrucipennis), Gallup. UTAH: Moab.

MEASUREMENTS: Length: 15-21 mm; width: 5.3

—7.6 mm.

TYPES. — Zopherus elegans Horn. Holotype (AN

SP 3920), Canon de Chelly, Arizona (not New Mexico

as stated by Horn).

Zopherodes otiosus Casey. The first of the two

specimens under this name in the Casey collection has

been selected and so labeled a lectotype by me. Both
NUMBE R 10 8 19

specimens also bear a USNM type number 46353 and

are from Fort Wingate, New Mexico.

Zopherodes verrucipennis Casey. Holotype (USNM

46354), Fort Wingate, New Mexico.

Zopherodes circumductus Casey. Holotype (USNM

46347), Fort Wingate, New Mexico.

Zopherodes woodgatei Casey. Holotype (USNM

46348), Jemez Springs, New Mexico.

SPECIMENS EXAMINED (in addition to the melanistic specimens listed above). — 41 from the localities

that follow. UNITED STATES. ARIZONA: Chinle,

Kayenta, Leupp, Marsh Pass, Tuba City. COLORADO:

Durango, Mesa Verde National Park, Montezuma

National Forest. NEW MEXICO: Albuquerque, Apache

National Forest, 11 miles west of Bernalillo, Bloomfield, Magdalena. UTAH : Moab, Torrey. Specimens

have been collected from 20 May to 29 August.

To clarify the above synonymy, it should be mentioned that both circumductus and woodgatei are

typical elegans in that they both have the whitebordered elytra and pronotum. Zopherodes otiosus

appears to be a melanistic phase of elegans, and verrucipennis was placed as a synonym of otiosus by Casey

himself (1911:254). Later (1924:305) Casey decided

that verrucipennis should be considered a variety of

otiosus, and the two were retained as separate entities

in Casey's collection according to L. L. Buchanan,

who transferred the collection from boxes to unit

trays.

Zopherus elegans Horn was reported from Durango,

Mexico, by Champion (1892:489, tab. 22 fig. 7).

These specimens are not elegans at all but rather

two undescribed species. These are described earlier

in this paper as solieri and championi.

19. Zopherus sanctaehelenae (Blaisdell),

new combination

PLATE 4: FIGURE 27

Zopherodes sanctaehelenae Blaisdell, 1931: 111.

This interesting species apparently represents an outlying isolated population derived from the same ancestral stock as was Z. granicollis. Because of its disjunctive distribution and constant morphological

characteristics, I prefer to regard it as a distinct

species.

Blaisdell described this species in great detail from

30 specimens, and the few additional specimens collected since then do not modify his description in any

way. Z. sanctaehelenae may be distinguished from all

other North American species of the genus except

opacus and the more strongly sculptured specimens of

gracilis by the very fine, widely spaced elytral tubercles.

From the latter two species it may be separated by the

more robust body proportions.

MEASUREMENTS. — Length 13.0-21.0 mm; width

5.5-8.0 mm.

TYPES.—Holotype, male (CAS 2986) and allotype, female (CAS 2987), Mount Saint Helena, Napa

County, California, 15 March (not July, as stated by

Blaisdell) 1930, A. T. McClay, T. W. Serins, E. G.

Linsley.

SPECIMENS EXAMINED. — 23 (including 10 paratypes labeled same as holotype and allotype).

UNITED STATES. CALIFORNIA: Mount Saint Helena and Pope Valley, both in Napa County. Specimens have been taken from 22 February to 9 May.

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Societe Entomologique de France, series 2, 2:287-

313.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Casey, Thomas L.

1907a. Notes on Chalcolepidius and the Zopherini. Canadian Entomologist, 39:29-46.

1907b. A Revision of the American Components of the

Tenebrionid Subfamily Tentyriinae. Proceedings

of the Washington Academy of Sciences, 9:275-

522.

1911. Notes on the Coccinellidae with Some General

Remarks and Synonymy. Memoirs on the Coleoptera, 2:246-259.

1924. Additions to the Known Coleoptera of North

America. Memoirs on the Coleoptera, 11:1—347.

Champion, George C.

1884. Biologia Centrali-Americana, Insecta, Coleoptera

(Tenebrionidae), 4(1) : 1-88, illustrated.

1886. Biologia Centrali-Americana, Insecta, Coleoptera

(Tenebrionidae), 4( 1): 137-264, illustrated.

1892. Biologia Centrali-Americana, Insecta, Coleoptera.

(Supplement to Heteromera), 4(1) :477-524, illustrated.

Crowson, Roy A.

1955. The Natural Classification of the Families of

Coleoptera. 187 pages. London.

Gebien, Hans

1910. Coleopterorum Catalogus, pars 15, Tenebrionidae,

XVIII, (I): 166 pages.

Gray, George R.

1832. Notices of New Genera and Species. In Griffith

and Pidgeon, The Animal Kingdom Arranged in

Conformity with Its Organization by the Baron

Cuvier. Volume 15 (Insecta, volume 2). 796

pages. London.

Guerin-Meneville, Felix E.

1844. Description de quelques Coleopteres de la NouvelleGrenade. Revue Zoologique, par la Sociite Cuvierienne, 7:8-19.

Haldeman, Samuel S.

1852. Insects, in Stansbury, Exploration and Survey of

t h e Valle y of th e Grea t Sal t Lak e of Uta h ... .

Pages 366-378, illustrated. Washington.

Hope, Frederic W.

1840. The Coleopterist's Manual, Part the Third, Containing Various Families, Genera, and Species, of

Beetles, Recorded by Linneus and Fabricius. Also,

Descriptions of Newly Discovered and Unpublished Insects. 191 pages, illustrated. London.

Horn, George H.

1867. Notes on the Zopheri of the United States. Transactions of the American Entomological Society,

1:159-162.

1870. Revision of the Tenebrionidae of America, North

of Mexico. Transactions of the American Philosophical Society, series 2, 14:253-404, illustrated.

1885. Contributions to the Coleopterology of the United

States. (Number 4). Transactions of the American

Entomological Society, 12:128-162, illustrated.

1894. The Coleoptera of Baja California. Proceedings of

the California Academy of Sciences, series 2, 4:302-

499, illustrated.

Lacordaire, Jean T.

1859. Genera des coleopteres . . . .Volume 5. 750 pages,

illustrated. Paris.

LaPorte, Francois L. N. de

1840. Histoire naturelle des animaux articulis, 1:324

pages, 2:564 pages. Paris.

LeConte, John L.

1851. Descriptions of New Species of Coleoptera, from

California. Annals of the Lyceum of Natural History of New York, 5:125-184.

Lucas, Pierre H.

1852. Note on Zopherus moreletii. Bulletin de la Sociite Entomologique de France: xxiii.

Salle, August

1849. Coleopteres nouveaux de l'Amerique (l r e partie).

Annales de la Sociite Entomologique de France,

series 2, 7:297-303.

Solier, Antoine J. J.

1841. Essai sur les Collapterides (suite). Annales de la

Sociite Entomologique de France, 10:29—51, illustrated.

Sturm, Jacob

1843. Catalog der Kafer-Sammlung von Jacob Sturm.

386 pages, 6 plates. Niirnberg.

Watt, J. Charles

1967. A Review of Classifications of Tenebrionidae

(Coleoptera). The Entomologist's Monthly Magazine, 102:80-86.
NUMBER 10 8 21

PLATE 1.—Genus Zopherus: 1, Z. laevicollis Solier, elytral apex; 2, Z. angulicollis Champion,

elytral apex; 3, Z. laevicollis Solier, ventral aspect of abdomen showing trilobed callus of terminal

sternum; 4, Z. mexicanus Gray, ventral aspect of abdomen showing arcuate callus of terminal

sternum; 5, Z. nodulosus haldemani Horn, ventrolateral aspect of prothorax showing appendiculate apex of antennal fossa; 6, Z. xestus new species, ventrolateral aspect of prothorax

showing rounded apex of antennal fossa; 7, Z. nodulosus haldemani Horn, antenna; 8, Z.

chilensis Gray, aedeagus (length: 6.8 mm.)-
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

11

PLATE 2.—Genus Zopherus: 9, Z. chilensis Gray, El Salvador (length: 42.5 mm); 10, Z.

nervosus Solier, Guerrero, Mexico (length: 24.6 mm); 11, Z. nodulosus nodulosus Solier, Sonora,

Mexico (length: 21.7 mm); 12, Z. nodulosus haldemani Horn, Bastrop, Texas (length: 23.1

mm); 13, Z. jourdani Salle, Tegucigalpa, Honduras (length: 25.8 mm); 14, Z. mexicanus Gray,

Hidalgo, Mexico (length: 19.3 mm).
NUMBER 10 8 23

17

21

19

PLATE 3.—Genus Zopherus: 15, Z. angulicollis Champion, Chihuahua, Mexico (length: 18.7

mm); 16, Z. laevicollis Solier, Chapingo, Mexico (length: 19.9 mm.); 17, Z. xestus, new

species, Chisos Mountains, Texas (length: 20.4 mm); 18, Z. championi, new species, Sanderson,

Texas (length: 14.3 mm); 19, Z. tristis LeConte, Baja California, Sur, Mexico (length: 20.0

mm); 20, Z. concolor LeConte. Bandelier National Monument, New Mexico (length: 19.8 mm);

21, Z. gracilis Horn, Ruby, Arizona (length: 18.2 mm).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

22 23 24

26 27

PLATE 4.—Genus Zopherus: 22, Z. uteanus (Casey), Utah (length: 16.2 mm) ; 23, Z. granicollis

granicollis Horn, Berkeley, California (length: 19.0 mm); 24, Z. granicollis ventriosus (Casey),

Sugar Pine, California (length: 17.3 mm); 25, Z. opacus Horn, Nevada (length: 17.2 mm);

26, Z. elegans Horn, Sandoval County, New Mexico (length: 18.8 mm) ; 27, Z. sanctaehelenae

(Blaisdell), Mount Saint Helena, California (length: 17.9 mm).

US. GOVERNMENT PRINTING OFFICE: 1972 4 84-312/4

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