CHARLES A. TRIPLEHO A Review of the Qenus
_ Zopherus of the World
(Coleoptera: i
Tenebrionidae)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 108
S E R I A L P U B L I C A T I O N S OF THE S M I T H S O N I A N INSTITUTION
The emphasis upon publications as a means of diffusing knowledge was expressed
by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti-
tution/Joseph Henry articulated a program that included the following statement:
"It is proposed to publish a series of reports, giving an account of the new discoveries
in science, and of the changes made from year to year in all branches of knowledge."
This keynote of basic research has been adhered to over the years in the issuance
of thousands of titles in serial publications under the Smithsonian imprint, com-
mencing with Smithsonian Contributions to Knowledge in 1848 and continuing
with the following active series:
Smithsonian Annals of Flight
Smithsonian Contributions to Anthropology
Smithsonian Contributions to Astrophysics
Smithsonian Contributions to Botany
Smithsonian Contributions to the Earth Sciences
Smithsonian Contributions to Paleobiology
Smithsonian Contributions to Zoology
Smithsonian Studies in History and Technology
In these series, the Institution publishes original articles and monographs dealing
with the research and collections of its several museums and offices and of profes-
sional colleagues at other institutions of learning. These papers report newly acquired
facts, synoptic interpretations of data, or original theory in specialized fields. These
publications are distributed by mailing lists to libraries, laboratories, and other in-
terested institutions and specialists throughout the world. Individual copies may be
obtained from the Smithsonian Institution Press as long as stocks are available.
S. DILLON RIPLEY
Secretary
Smithsonian Institution
SMITHSONIAN CONTRIBUTIONS TO
ZOOLOGY
NUMBER 108
Charles A. Triplehorn A R e v i e w o f t h 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 in-
habit 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.
UNITED STATES GOVERNMENT PRINTING OFFICE
WASHINGTON : 1972
For sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, D.C. 20402 - Price 45 cents {paper cover)
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 interest-
ing group of beetles was by Casey (1907a). In his
paper he divided the genus Zopherus into four gen-
era, 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 es-
tablished, determinations at the species level were
uncertain.
Blaisdell (1931) presented a key to the California
species of Zopherodes from which he omitted Z. grani-
Charles 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 "Zo-
phorus chilensis, the type of a new genus near Zopho-
sis." 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 emend-
ing Gray's spelling and therefore should not be cited
as the author of the genus as many subsequent au-
thors and catalogers have done.
Solier (1841:30) placed Zopherus in his tribe Zo-
pherites, along with Nosoderma Dejean and Dicero-
deres 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 slight-
ly (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 de-
scribed. The genus was thus well known and very
stable until Casey (1907a) divided it into four gen-
era. He designated a type species for each as follows:
Megazopherus containing only chilenis Gray; Zo-
pherodes with tristis LeConte as type species; Zophe-
rinus with limbatus Casey as type species, and Zophe-
rus 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 con-
clusions 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 argu-
ments for a separate family and I concur with Arnett
(1963:662), who places the group in the tribe Zoph-
erini, subfamily Asidinae.
Morphology
All members of the genus Zopherus are elongate and
subcylindrical, with extremely thick and hard integu-
ments. 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 pro-
notum are easily scored with a sharp instrument, the
integument having the physical properties of an ex-
tremely 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 pat-
terned black and white species to become uniformly
shiny black. Surface sculptures (punctures and tu-
bercles) 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 in-
dicated 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-
N U M B E R 1 0 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 incon-
spicuous, usually hidden beneath the base of the pro-
notum when the body is in repose.
ELYTRA. ? The elytra are fused but the sutural line
is always apparent. The apices are always conspicuous-
ly 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 pre-
sents 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 ex-
tremely fine and compact ridges of golden hairs that
extend from base to apex of each extremity, and con-
spicuously 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. Fe-
males tend to be somewhat larger and stouter than
males, but body proportions are, in general, unre-
liable 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 Zo-
pherus. 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 vari-
ations.
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 describ-
ing 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 speci-
mens upon which this report is based came from
numerous sources, and I am indebted to many in-
stitutions and individuals for lending their collec-
tions 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. Cham-
pion material, which proved invaluable in working
out the Central American species. Through the cour-
tesy 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 count-
less 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. Triple-
horn. 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-Pen-
rose Fund) and 1967 (Grant No. 4597-Penrose
Fund). Grateful acknowledgment is made to this so-
ciety 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. Her-
man, Jr.
ANSP Academy of Natural Sciences of Philadel-
phia: Harold J. Grant, Jr.
ASU Arizona State University, Tempe: Frank F.
Hasbrouck and Stanley C. Williams.
BMNH British Museum (Natural History), Lon-
don: C. M. F. von Hayek and J. Balfour-
Browne.
N U M B E R 1 0 8
BYU Brigham Young University, Provo, Utah:
Vasco M. Tanner.
CAS California Academy of Sciences, San Fran-
cisco: Hugh B. Leech.
CAT Charles A. Triplehorn private collection.
CMP Carnegie Museum, Pittsburgh: George Wal-
lace.
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, Bra-
zil : 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. San-
derson.
INIA Instituto Nacional de Investigaciones Ag-
ricolas, Chapingo, Mexico: Francisco Pa-
checo 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 Univer-
sity, 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: Ev-
erett 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, Washing-
ton, 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 scat-
tered 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 nar-
rower than elytra 20
Clypeal punctures small, shallow and widely separated; species of slender proportions,
pronotum usually as wide as or wider than elytra 21
N U M B E R 1 0 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 recog-
nized 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
Tec la . G U A T E M A L A : Asuncion Mi ta , J u t i a p a .
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 con-
spicuously 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, repre-
senting 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, marmora-
tus), 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 Guer-
rero; 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.?Halde-
man, 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 re-
spects 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, includ-
ing 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 pro-
nounced 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 Guer-
rero; 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 Tehua-
can 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 con-
stant 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.
N U M B E R 1 0 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, Brown-
wood, 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 Braun-
fels, 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 south-
west of Cuencame.
I have seen specimens labeled "Guatemala," "Cali-
fornia," "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 speci-
mens 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 Trini-
taria (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 sum-
marized 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 syn-
onymy 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 con-
siderably 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 Hi-
dalgo (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 inter-
mediate 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 color-
ation, 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 speci-
mens 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
N U M B E R 1 0 8 11
distinguished from all other species of Zopherus except
xestus by the peculiar formation of the ventral cal-
losity of the terminal abdominal sternum. In both
laevicollis and xestus there is a prominent ridge pro-
jecting cephalad from the normal bilobed callosity,
causing this structure to appear trilobed as described by
Solier.
After studying a good series of Champion's speci-
mens of venosus and specimens which he determined
as laevicollis, I concur with his earlier (1884) con-
viction 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: Maza-
mitla. 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 High-
way, 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 Chalchi-
comula (Puebla), Juquila and Calpulalpan (Oaxa-
ca ). Specimens have been taken in every month
except March.
9. Zopherus xestus, new species
PLATE 3: FIGURE 17
DESCRIPTION.?Holotype, sex undetermined. Elon-
gate, 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 punc-
tures 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 prom-
inence 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 pro-
sternum; 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 form-
ing 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 nu-
merous 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 sculptur-
ing. 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 Coun-
ty, 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. Hafer-
nik (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 prom-
inent, 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 round-
ed 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. Ely-
tra 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 lat-
eral 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 fre-
quently are contiguous, inflexed portion entirely black,
finely and sparsely punctured. Mesosternum and meta-
sternum 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 punc-
tured.
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 fos-
sae 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 para-
types, all from Villa Lerdo, Durango, Mexico, col-
lected by Hoge (BMNH); one paratype, same data
N U M B E R 1 0 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 emargi-
nate, dorsal surface coarsely but shallowly and sparsely
punctured, each puncture with a short, coarse, yellow-
ish seta, ventral surface smooth except for a few wide-
ly spaced punctures laterally; mentum broadly trans-
verse, 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; an-
tennal fossae rounded apical ly, prosternum coarse-
ly, 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 bor-
ders white, irregularly spotted with black blotches, in-
flexed portion entirely black, coarsely and sparsely
punctured. Mesosternum and metasternum coarsely
and irregularly punctured; abdominal sterna coarse-
ly and densely punctured medially but practically
impunctate laterally; ventral callus of terminal abdom-
inal 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 para-
types 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. Schaff-
ner (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 recog-
nize, but one of the most difficult to characterize. The
body is subcylindrical, greatly thickened dorsoventral-
ly, with the elytral margins parallel throughout a
greater portion of their length and abruptly constrict-
ed 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 re-
stored by soaking the specimen in ether. It appears
to be of no taxonomic significance.
Casey (190 7b: 467) considered regarding the spe-
cies 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 ex-
hibited 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." Zo-
pherodes 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 vari-
able 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, Ba-
boquivari Mountains, Buckeye, Bumble Bee, Chandler,
Florence, Granite Reef, Mesa, Nogales, Organ Pipe
Cactus National Monument, Parker, Phoenix, Pres-
cott, Riverside, Safford, San Carlos Lake, Santa
Catalina Mountains, Tacna, Tempe, Tucson, Vail,
Wellton, 12 miles east of Wenden, Wickenburg,
Yuma. CALIFORNIA: Blythe, Borrego, Calexico, Cali-
ente, Coachella Valley, Carrizo Springs, Holtville, Im-
perial, 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 fre-
quently form transverse wavy plicae. The pronotal
punctures are sparse and simple, at least never forming
rough granules, and the prosternum is usually rough-
ly sculptured with punctures rather than raised tuber-
cles. 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 con-
color.
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, repre-
sent this species in the Casey collection. The first speci-
men 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: Albu-
querque, Bandelier National Monument, Bent, Cloud-
croft, Conchas Dam, Coolidge, Corona, Duran, High
Rolls, Hot Springs, Jemez Springs, Las Vegas, May-
hill, Mescalera Reservation, Omega, Romeroville,
Ruidoso, Santa Fe, Tecolote. TEXAS : Davis Moun-
tains, 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
N U M B E R 1 0 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. Speci-
mens 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 un-
forgivable 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 ob-
vious 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 mar-
gins 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 ab-
dominal segment is broadly arcuate, with an acutely
rounded and prominent median lobe directed ceph-
alad. 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 collec-
tion. It is rather typical of those found in the Flag-
staff 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 repre-
sented 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 syn-
onymyzed 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 se-
lected 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 col-
lection 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 speci-
mens 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 vari-
ous 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, Gali-
uro 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. N E W MEX-
ICO: 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 ven-
triosus in elytral sculpture but may be distinguished
by the fine, sparse punctation of the head. In grani-
collis 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 some-
times they are smaller and more distantly spaced, ap-
proaching the situation in opacus. It is this latter
variation that Casey named mormon. Enough inter-
mediate forms have been studied to convince me that
only one species is involved. In all specimens examined
the pronotum is coarsely and densely granulate-punc-
tate.
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 col-
lection 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). U T A H : 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 col-
lected 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
N U M B E R 1 0 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 moun-
tains of southern California, and northern Baja Cali-
fornia. 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 some-
what 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 elon-
gated as in typical ventriosus. I have seen this same
condition in series in which most individuals were
coarsely tuberculate and had assumed that it repre-
sented 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 in-
dicating "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. grani-
collis.
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 arrange-
ment 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 Monu-
ment, 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, Ru-
morosa, 2 miles south of El Toro (Sierra Juarez).
Specimens have been collected in every month ex-
cept 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 punc-
tures 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 sub-
species, granicollis and ventriosus, apparently occupy-
ing 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 Mead-
ow, 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 Mead-
ow 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 trans-
versely elongate, with thin linear crests and a bit
closer together than usual. Horn had a similar speci-
men from Nevada, so I assume that this can be re-
garded 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 ven-
triosus. 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 identi-
fied by the whitish lateral borders of the pronotum
and elytra. The pronotum is always coarsely granulate-
punctate, 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 re-
lief. 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 melan-
istic 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 Moun-
tains, White Mountains, 8-15 miles northeast of
Whiteriver. N E W MEXICO: Apache National Forest,
Fort Wingate ('ectotype and paratype of otiosus and
holotype of verrucipennis), Gallup. U T A H : 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
N U M B E R 1 0 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 melanis-
tic 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, Bloom-
field, Magdalena. UTAH : Moab, Torrey. Specimens
have been collected from 20 May to 29 August.
To clarify the above synonymy, it should be men-
tioned that both circumductus and woodgatei are
typical elegans in that they both have the white-
bordered elytra and pronotum. Zopherodes otiosus
appears to be a melanistic phase of elegans, and ver-
rucipennis 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 out-
lying isolated population derived from the same an-
cestral stock as was Z. granicollis. Because of its dis-
junctive 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 col-
lected 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 allo-
type, 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 para-
types labeled same as holotype and allotype).
UNITED STATES. CALIFORNIA: Mount Saint Hel-
ena and Pope Valley, both in Napa County. Speci-
mens have been taken from 22 February to 9 May.
Literature Cited
Arnett, Ross H., Jr.
1963. The Beetles of the United States. 1112 pages, illus-
trated. Washington, D.C.: The Catholic Univer-
sity of America Press.
Blaisdell, Frank E.
1923. Expedition of the California Academy of Sciences
to the Gulf of California in 1921. The Tene-
brionidae. Proceedings of the California Academy
of Sciences, series 4, 12:201-288.
1931. A New Species of Zopherodes from Central Cali-
fornia (Coleoptera: Tenebrionidae). Pan-Pacific
Entomologist, 7 (3) : 111-114.
Blanchard, Charles E.
1861. In D'Orbigny, Dictionnaire universel d'histoire
naturelle. Atlas, 2. Paris.
Boddy, Dennis W.
1965. Family Zopheridae. In Hatch, The Beetles of the
Pacific Northwest. Part IV: 77-79, illustrated.
Seattle: University of Washington Press.
Boving, Adam G., and Frank C. Craighead
1931. An Illustrated Synopsis of the Principal Larval
Forms of the Order Coleoptera. Entomologia
Americana, 11:1-351, 125 plates.
Brcme, Francois de
1844. Insectes Coleopteres nouveaux ou peu connus.
Premiere et deuxieme decades. Annales de la
Societe Entomologique de France, series 2, 2:287-
313.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Casey, Thomas L.
1907a. Notes on Chalcolepidius and the Zopherini. Cana-
dian 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 Coleop-
tera, 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, il-
lustrated.
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 Nouvelle-
Grenade. Revue Zoologique, par la Sociite Cuvi-
erienne, 7:8-19.
Haldeman, Samuel S.
1852. Insects, in Stansbury, Exploration and Survey of
t h e V a l l e y of t h e G r e a t S a l t L a k e of U t a h . . . .
Pages 366-378, illustrated. Washington.
Hope, Frederic W.
1840. The Coleopterist's Manual, Part the Third, Con-
taining Various Families, Genera, and Species, of
Beetles, Recorded by Linneus and Fabricius. Also,
Descriptions of Newly Discovered and Unpub-
lished Insects. 191 pages, illustrated. London.
Horn, George H.
1867. Notes on the Zopheri of the United States. Trans-
actions of the American Entomological Society,
1:159-162.
1870. Revision of the Tenebrionidae of America, North
of Mexico. Transactions of the American Philo-
sophical 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 His-
tory of New York, 5:125-184.
Lucas, Pierre H.
1852. Note on Zopherus moreletii. Bulletin de la So-
ciite 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, illus-
trated.
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 Mag-
azine, 102:80-86.
N U M B E R 1 0 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 appen-
diculate 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).
N U M B E R 1 0 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

Publication in Smithsonian Contributions to Zoology
Manuscripts for serial publications are accepted by the Smithsonian Institution Press, sub-
ject to substantive review, only -through departments of the various Smithsonian museums. Non-
Smithsonian authors should address inquiries to the appropriate department. If submission is
invited, the following format requirements of the Press will govern the preparation of copy.
Copy must be typewritten, double-spaced, on one side of standard white bond paper, with
1%" top and left margins, submitted in ribbon copy with a carbon or duplicate, and accompa-
nied by the original artwork. Duplicate copies of all material, including illustrations, should be
retained by the author. There may be several paragraphs to a page, but each, page should begin
with a new paragraph. Number consecutively all pages, including title page, abstract, text, litera-
ture cited, legends, and tables. The minimum length is 30 pages, including typescript and illus-
trations.
The title should be complete and clear for easy indexing by abstracting services. Taxonomic
titles will carry a final line indicating the higher categories to which the taxpn is referable:
"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify
the author on the first page of text with an unnumbered footnote that includes his professional
mailing address. A table of contents is optional. An index, if required, may be supplied by the
author when he returns page proof.
Two headings are used: (1) text heads (boldface in print) for major sections and chap-
ters and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further
headings may be worked out with the editor.
In taxonomic keys, number only the first jtem of each couplet; if there is only one couplet,
omit the number. For easy reference, number also the taxa and their corresponding headings
throughout the text; do not incorporate page references in the key.
In synonymy, use the short form (taxon, author, date:page) with a full reference at the
end of the paper under "Literature Cited." Begin each taxon at the left margin with subse-
quent lines indented about three spaces. Within an entry, use a period-dash (.?) to separate
each reference. Enclose with square brackets any annotation in, or at the end of, the entry.
For references within the text, use the author-date system: "(Jones 1910)" and "Jones
(1910)." If the reference is expanded, abbreviate the data: "Jones (1910:122, pi. 20: fig. 1)."
Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they
should not contain rules. Formal tables must be submitted as pages separate from the text, and
each table, no matter how large, should be pasted up as a single sheet of copy.
Use the metric system instead of, or in addition to, the English system.
Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed
throughout the printed text. They will be termed Figures and should be numbered consecu-
tively; however, if a group of figures is treated as a single figure, the components should be
indicated by lowercase italic letters on the illustration, in the legend, and in text references:
"Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as
full pages on a different stock of paper, they will be termed Plates, and individual components
should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2)* Never combine the
numbering system of text illustrations with that of plate illustrations. Submit all legends on
pages separate from the text and not attached to the artwork. An instruction booklet for the
preparation of illustrations is available from the Press on request
In the bibliography (usually called "Literature Cited"), spell out book, journal, and
article titles, using initial caps with all words except minor terms such as "and, of, the." For
capitalization of titles in foreign languages, follow the national, practice of each language.
Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume,
number, and page citations: "10(2) :5-9." Spell out such words as "figures," "plates," "pages."
For free copies of his own paper, a Smithsonian author should indicate his requirements
on "Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will
receive 50 free copies; order forms for quantities above this amount with instructions for pay-
ment will be supplied when page proof is forwarded.