Memoirs of the American Entomological Society

MEMOIRS
OF THE

AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 26

A TAXONOMIC REVISION
OF THE AQUATIC BEETLE GENUS
LACCOPHILUS (DYTISCIDAE)
OF NORTH AMERICA

BY

JAMES R. ZIMMERMAN

PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA

1970

3

MEMOIRS
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MVGEwherAUNe EEN cOMOLOGCICAL SOCIETY
NUMBER 26

A TAXONOMIC REVISION
OF THE AQUATIC BEETLE GENUS
LACCOPHILUS (DYTISCIDAE)
OF NORTH AMERICA

BY

JAMES R. ZIMMERMAN

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PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES

PHILADELPHIA

1970

SELWYN S. ROBACK
EDITOR

(Issued August 18, 1970)

PRINTED IN THE UNITED STATES OF AMERICA

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MEM. AMER. ENT. SOC., 26

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MEMOIRS
OF THE

AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 26

A TAXONOMIC REVISION OF THE AQUATIC BEETLE
GENUS LACCOPHILUS (DYTISCIDAE)
OF NORTH AMERICA*

BY
JAMES R. ZIMMERMAN

Department of Biology
New Mexico State University
Las Cruces, New Mexico 88001

INTRODUCTION

The aquatic beetle genus Laccophilus occurs on all continents ex-
cept Antarctica — as well as on many nearby islands and island chains.
Zimmermann (1920) included 167 species of Laccophilus in the world
list. Of this number 27 are recorded from North America and the
West Indies. Most of the remainder are in the tropics. Later lists
(Leng 1920, 1927, 1933; Blackwelder 1939, 1944; Blackwelder and
Blackwelder 1948) increase the total to 32. In North America there
is a good representation of the genus, but not an extremely heavy
concentration of species.

Several experienced workers in the Dytiscidae have commented
on the difficulty in identifying and discriminating the species (Sharp
1882a, p. 287; F. Balfour-Browne 1940, p. 180; Leech 1948b; J.
Omer-Cooper 1965). This problem may have been one important
reason why no extended revision has been attempted in North Amer-
ica. The last revision (Crotch 1873) was not done in a modern
sense, but was a summation of literature and descriptions of new

1 This study was supported by NSF grants G17936 and G23585.

MEM. AMER. ENT. SOC., 26

2 THE GENUS LACCOPHILUS

species. Several regional treatments were done in this century as part
of faunal studies — Blatchley in Indiana and Florida (1910, 1919),
Young in Florida (1954), Leech in Baja California (1948), Hatch
in the Pacific Northwest (1953), and Anderson in Utah (1962).
They were not intended to be revisionary; but all of them, especially
Leech and Young, stress the need for such a study. It was evident
that a revision of at least continental scope would be required to de-
fine species limits and for an understanding of the evolutionary diver-
gence taking place in North American Laccophilus.

This study is a revision of the adults of North American species.
Species with most of their ranges included in the Nearctic Realm are
treated more thoroughly than primarily Neotropical ones which have
extended their ranges into North America. West Indian species are
generally omitted.

Determination of species limits requires studies of population vari-
ation. Some of the difficulties encountered by earlier workers were
due to the lack of knowledge of the limits of variation. The diffi-
culties in identification of Laccophilus species are minimal once that
information is available.

Since a generic study also can provide information about the
amount of differentiation in the formation of species, considerable
attention was given to the recognition of races as well as species. In
this respect there was close agreement in Laccophilus with the study
of Burns (1964) on the skipper genus Erynnis. In his revision he
states, “Locality data accumulated from accurately determined speci-
mens show that much of the genus falls into complexes of geographi-
cally complementary forms. Each complex is broadly distributed in
North America, and each is composed of entities that are both mor-
phologically and biologically one another’s near relatives. Some of
these complexes may almost equally well be treated as polytypic spe-
cies or as superspecies. Usually the choice is arbitrary and subjective,
so that classification only reflects — imperfectly — the varying degrees
of relationship that exist among the populations in nature.”

Lumping of taxa has been the guideline in this study of Lac-
cophilus. Allopatric groups previously considered species have been
reduced to subspecies if it was felt there was enough similarity (obvi-
ously a subjective judgment). The same guideline applies to new
forms described as subspecies. The presence of intermediates be-

JAMES R. ZIMMERMAN 3

tween two morphologically different and geographically replacing
populations was considered to be evidence of intergradation; and the
populations were thus given subspecific rank. If, however, there was
a test of sympatry with a consistent difference and no intermediates,
then specific rank had to be accorded, regardless of similarity. Some
species are more similar in appearance than are some races within the
same species (for example, L. sonorensis, L. q. quadrilineatus, and
L. q. tehuanensis ).

I wish to acknowledge the following individuals and museums for
making material available for study: Patricia Vaurie, Jerome Rozen,
American Museum of Natural History (AMNH); Floyd Werner, Uni-
versity of Arizona Museum of Entomology (ARI); J. Balfour-Browne,
British Museum (Natural History) (BM); Jerry A. Powell, Paul
Hurd, University of California, Berkeley (BERK); Richard Bohart,
A. T. McClay, University of California, Davis (DAV); Ross H. Ar-
nett, Eileen Van Tassell, Catholic University of America (CUA);
George Wallace, Carnegie Museum (CNG); Henry Dietrich, Cornell
University (CNL); James Sublette, Eastern New Mexico University;
Rupert Wenzel, Field Museum of Natural History (FM); P. J. Darl-
ington, Jr.; Harvard University Museum of Comparative Zoology
(MCZ); Jean L. Laffoon, Iowa State College (ISU); G. Fogel, A.
Capart, Institut Royal des Sciences Naturelles de Belgique, Brussels;
Fred S. Truxal, Los Angeles County Museum (LACM); Guy Colas,
Museum National d’Histoire Naturelle, Paris; Ronald B. Willson, Ro-
land L. Fischer, Michigan State University (MCHS); John D. Lattin,
Oregon State University (ORES); H. Radcliffe Roberts, Academy of
Natural Sciences of Philadelphia (ANSP); Horace Burke, Joseph C.
Shafiner, Texaxs A & M University (TAM); Russel D. Anderson
(RDA); J. L. Carr (CARR); Burris McDaniels; Joe Schuh (SCH).

In addition, the following people have collected specimens for me:
Robert D. Ohmart, Robert L. Smith, Kenneth L. McWilliams, and
Robert N. Gennaro.

Georgianna Gunaji and William Doersam, National Science Foun-
dation undergraduate research participants, also contributed directly
to some of the information in this paper. Mr. Doersam measured
most of the specimens of L. pictus and enumerated the locality records
for that species. Mrs. Gunaji helped curate many specimens and
assembled numerous locality records.

MEM. AMER. ENT. SOC., 26

4 THE GENUS LACCOPHILUS

Mr. Anthony Smith has been especially helpful. He has collected
hundreds of specimens and has written the computer program used in
the analysis of the statistical data.

Finally, special thanks are due to Frank N. Young, Indiana Uni-
versity and University of Michigan Museum of Zoology (UMMZ),
Hugh Leech, California Academy of Sciences (CAS), and Paul
Spangler, United States National Museum (USNM), for the advice
and encouragement as well as the specimens they have contributed
during this study.

MATERIALS AND METHODS

Collecting. — A large number of the specimens from Indiana,
Kansas, and Oklahoma, the Southwest, and Mexico, were collected by
the writer. Sweeping the water with a metal sieve or strainer was the
most common method. An eight inch diameter wire strainer which
can be found at the housewares section of grocery and hardware stores
was modified and strengthened by replacing the handle with another
heavier one on which was fitted a strong metal ring of approximately
the same diameter. The commercial strainers have two projecting
wire loops on the front that can be bent down over the heavy ring to
help hold the wire sieve in place. The wire that formerly fit into the
wooden or plastic handle that came with the strainer is inserted into
the heavy wooden handle. A household strainer so modified will last
several months or even years. The sieve can easily be replaced if the
wires break. Short sweeps through water of a few inches depth, with
the rim of the strainer just grazing the bottom of the pond or stream,
yields the best results. Particular attention is given to grass, sedges,
and other vegetation around the margins, and to debris or gravel on
the bottom. Almost any freshwater habitat may harbor Laccophilus.
Temporary roadside ditches, quarry pools, cattle tanks or ponds, small
streams, margins of lakes, seepage areas, and sloughs have been suc-
cessful collecting localities. Usually large river margins, riffles, water
of more than one foot deep, and hot springs do not provide favorable
habitat for the members of this genus. Young’s (1954) statement
about the distribution of water beetle populations is applicable at this
point. “The actual factors which influence the maintenance of a pop-
ulation of water beetles are very numerous. The presence of food
and pressure from predators are two important factors which vary

JAMES R. ZIMMERMAN 5

greatly with local conditions, and together with the need for replen-
ishing the air in the tracheal system and a terrestrial situation for
pupation, confine many species throughout their lives to a narrow
shore-line strip. Water beetle populations are thus often not directly
correlated with the volume of water, but with the extent of suitable
shore line.”

Once taken, each specimen is picked from the sieve with a feather-
tip forceps and placed in a vial of 70 per cent ethanol and a few drops
of glycerin. Vials were fitted with screw top plastic lids which had
polyethylene cone liners to stop evaporation. Locality data included
distance to nearest recognizable village, town, or city, and county (if
in the United States), kind of habitat (stream, pond, roadside ditch),
amount of emergent and submergent vegetation, nature of substrate,
condition of water, surrounding plant life form, date, and collector(s).
Elevation was frequently, but not always, included. There was an
attempt to take all species of water beetles encountered, but naturally
other genera would be less well represented in the collections.

Preparation and Examination. — Water beetles store fats and oils,
and these can diffuse through breaks in the dried cuticle. Eventually
a coating of grease covers most of the insects. The grease and the
dust it collects obscure colors and structures. Hence, it is necessary
to degrease specimens — both those recently collected and those that
have been mounted for some time. The most successful method has
been to soak them in ethyl ether. The insects are left in the ether
until there is no increase in the yellow-orange color that results from
dissolving the grease. This period may last from two hours to two
days, but rarely longer. Once degreased, the specimens may be
brushed clean with a fine brush dipped in 70 percent alcohol. Ap-
pendages are less brittle and less apt to break once the insects are
cleaned. Insufficiently degreased specimens frequently come unglued
from points.

Male genitalia must be extracted for examination. Fresh alco-
holic specimens permit extraction with no further treatment, but dried
ones must be softened. This can be done by gently boiling the beetles
in water for a few minutes. Extraction is done under a binocular dis-
secting microscope by holding the insect with a pair of forceps (trial
and error will show which kind is best) in one hand and grasping the
aedeagus with sharpened watchmaker’s forceps in the other. It is

MEM. AMER. ENT. SOC., 26

6 THE GENUS LACCOPHILUS

better if the aedeagus and parameres are not detached from the insect;
but, if they are, they can be glued to a paper point and then placed
on the same pin as the insect. After drying, it is best to place the
specimen into ether for an hour or two. Dry again before gluing on
a point. Shellac has proven to be the best glue, but white glue and
clear fingernail polish are also useful.

The ovipositor should also be exposed so that its form and number
of teeth can be determined. This can be done easily on fresh or alco-
holic specimens with much the same method used for male genitalia,
but is usually more difficult in dried specimens that have been relaxed
by boiling.

Descriptive characters. — Each species description is introduced by
an assessment of general appearance which includes 1) size, 2) over-
all impression of color, 3) type of elytral pattern, 4) presence or ab-
sence of a metacoxal file, 5) length of prosternal process, and 6) form
of the ovipositor.

1. Size is rated as small, medium, or large. Small insects are three
to four millimeters long, medium ones are four to five millimeters,
and large ones are over five millimeters. Each description has the
extremes given for length and width.

2. The species are variously some degree of yellow, red, brown,
black, and their combinations. Megascopically, irrorated species dor-
sally appear some shade of brown; but others have strongly contrast-
ing patterns of reddish-brown and yellow, or black and yellow. The
method of killing and preserving has altered the colors in many speci-
mens; and this, along with the blending of colors, makes it difficult to
give precise color descriptions. It is better, perhaps, to give more
attention to whether a part of the exoskeleton is pale, usually some
shade of yellow, or darkened by the admixture of other colors.

3. Elytral patterns provide the best diagnostic character in Laccophi-
lus except for the male aedeagus. Patterns may be irrorated, uni-
colorous, suffused, marmorated or variegated. About twelve are of
the first type in which the pigment has been applied as individual
grains of sand. Each dot may be discreet or form a beadlike string,
or the pigment may spread sufficiently to make a continuous pattern
and thus appear suffused, marmorated, or variegated. The most com-
mon elytral color is some shade of brown, especially in the irrorated

JAMES R. ZIMMERMAN 7

species. Irroration is seldom uniform over the pattern, and there may
be continuous color — particularly on the margins and in the pos-
terior half of the elytra.

Although there is considerable individual variation in pattern,
there is frequently enough uniformity in a race or species to permit
positive identification of individuals regardless of sex, and in some
instances to allow one to draw conclusions about the presence or
absence of intergradation.

Elytral patterns help to break up the body outlines and in some
cases, especially in irrorated species, provide concealing coloration.
Irrorated species are extremely difficult to detect in water that has a
mud bottom, or debris, or that has even the slightest degree of tur-
bidity. Other species have strongly contrasting patterns (L. pictus
and L. horni, for example), and these most commonly occur in moun-
tain stream pools of the Southwest and Mexico which have clear water
and granitic gravel bottoms. The bright yellow markings on a dark
background make these animals virtually invisible until they move.
Young (1960) discusses his observations and conclusions on the im-
portance of color patterns in Laccophilus and other Southwestern and
Mexican genera of water beetles. He believes the patterns have a
strong selective advantage in the avoidance of predation.

4. An important secondary sexual character is the presence of a series
of ridges on the metacoxal plates on males of some species (a much
less well-developed file occurs on a few females). It is a stridulatory
file that is rubbed by the upper surface of the hind femur. The num-
ber and coarseness of the ridges vary among species and among the
races of species. A count is given for each species and race, but later-
ally the ridges often grade into the cuticular microreticulation and
makes the number given only an estimate. It is considered an im-
portant diagnostic character, but the character may have arisen in
different lines within the genus. The file is present in one race of
L. gentilis, but absent or only faintly suggested in the other. It is a
sound producing device that merits attention as the basis of a behav-
ioral adaptation which operates as a sexual isolation mechanism.

5. The presence of an acuminate, unispinose prosternal process lying
in the same plane as the prosternum is a generic feature of Laccophi-
lus. Sharp (1882a), without trying to assign taxonomic rank, placed

MEM. AMER. ENT. SOC., 26

8 THE GENUS LACCOPHILUS

a strong emphasis on the length of the process in his separation of
species. He distinguished between those with a long or a short process.
In the long one the process, and the groove on the metasternum which
receives it, extend posteriorly to an imaginary line drawn across the
hind margins of the mesocoxal cavities. Most North American Lac-
cophilus have a short process, however, in which neither tip nor
groove are that long. Some are intermediate and do not fit into either
category as reflected in Sharp’s descriptions (1882b) of L. duplex and
its synonym, L. optatus. He described the first as having a process of
moderate length and the second as having an elongate process. Since
the position of the process is variable due to the flexing of the pro-
thorax and since the length of the groove is sometimes difficult to
assess even with the best stereoscope and lighting, it is not surprising
that errors of interpretation result. It is probably one of the more
important phylogenetic characters in the genus, however.

6. There are distinctly different forms of the ovipositor in North
American Laccophilus. The first and most common type is one in
which the teeth are small, triangular, and tightly spaced, or sawlike
(fig. 42). It appears to be adapted for inserting eggs into tough tis-
sues by means of a sawing action. The other less common type (rake-
like) has the distal four or five teeth long and widespaced (fig. 188).
It must be used for oviposition in softer and weaker material than the
sawlike type. All species having the rakelike ovipositor are probably
monophyletic since they share so many other characters, but those
with a sawlike one appear to be composed of several different lineages.
Other characters. —
7. Microreticulation. F. Balfour-Browne (1940) maintains the only
basic sculptures in the cuticle of the Dytiscidae (and other Hydra-
dephaga) are reticulation and punctation. Laccophilus are covered
with a fine network of roughly polygonal cells impressed into the sur-
face. These cellules are frequently elongated on the metacoxal plates
and epipleura. The most striking modification for taxonomic use,
however, is that in some species a secondary mesh is formed by the
weakening of the primary network forming the individual cellules.
The tendency to form a stronger secondary network is termed “double”
and the lack of that tendency is “single” — following the terminology
of Young (1963). Balfour-Browne used elytra boiled in caustic

JAMES R. ZIMMERMAN 9

soda, permitting examination with transmitted light; but this is not
necessary for Laccophilus. They can be observed with a magnifica-
tion of 40 diameters or more in oblique reflected light if the specimen
is thoroughly cleaned.

Elytral microreticulation varies most among species, but there is
also some variation of the microreticulation on the head and prono-
tum. Males show the doubled condition more strongly than females.
As a general rule those species with a rakelike ovipositor have a single
condition, and species with a sawlike ovipositor have a double con-
dition. Exceptions are L. peregrinus for the former and L. biguttatus
for the latter.

Laccophilus species have almost no punctation on the elytra ex-
cept for the usual two or three faint rows of setigerous punctures
which seem common to all species with little noticeable variation.

8. Head. The head and its appendages are remarkably uniform in
North American Laccophilus, but the antennae of L. biguttatus are
slightly thickened when compared to other species. The only other
variable feature is a row of elongate punctures (supraclypeal seam)
that lie immediately above the clypeal margin. They may parallel the
margin, or arch upward from it, or diverge upward at the sides.

9. Pronotum. Even though Laccophilus species have highly similar
outlines, there is sufficient variation in shape and in the length of the
pronotum to make comparison of ratios reflecting these differences
worthwhile. Two values are given in each species description. The
first is the width of the pronotum at the head, divided by the width of
the pronotum at its base (WH/PW); and the second is the length of
the pronotum from the head to the apex where it meets the elytral
median suture, divided by the pronotal width (LP/PW).

The first value can be viewed in different ways. It shows a) how
sharply the body converges anteriorly, or b) how wide the head is
compared to the pronotum, or c) how ovate is the body. A high
value (for example, 0.73 in L. gentilis) indicates a wider head and a
more elongate body. Young (1954) notes that the narrow body form
is characteristic of gentilis, and this rational value then is a reflection
of that shape. On the other hand, L. maculosus has values of 0.66 to
0.68 and is obviously more ovate and more convergent anteriorly.

The second value reflects the acuteness of the angle formed by the

MEM. AMER. ENT. SOC., 26

10 THE GENUS LACCOPHILUS

hind margin of the pronotum at the elytral median suture. The higher
the value, the more acute is the angle (0.46 in L. pictus insignis); and
a lower value indicates a more obtuse angle (0.38 in L. horni).

The variation in the two ratios has not been tested statistically,
and the values given are approximate means. Even as approxima-
tions, they do give some indication of relationships. For example,
L. horni and L. leechi, which are similar in many other ways, have
identical values for both ratios. L. pictus, a species which is super-
ficially similar to L. horni, is quite different in shape of the pronotum
as revealed by the two values — suggesting less close relationship.

10. Epipleura. Considerable individual, racial, and specific variation
exists among females in the shape of the elytral epipleura. In their un-
modified condition as seen in males and many females, the epipleura
are widest anteriorly and narrows in a smooth curve to little more than
a thin line (fig. 2) at about the level of the first visible abdominal
sternite. In many females, however, the narrow posterior section is
expanded into a projecting lateral flange (figs. 276-278). Flanges
may never occur in a species (L. spangleri), or may occur in nearly
every female in some species (L. fuscipennis), or may be character-
istic of nearly every female in one subspecies and be rare in its sister
subspecies (L. maculosus and L. fasciatus). The degree of epipleural
expansion varies among individuals of a population — from barely
perceptible with 20 power magnification to easily observable mega-
scopically (L. maculosus shermani). In contrast the position of the
flange seems to be more characteristic of subspecies and species and
is not an individual variation. The best example is the situation in
L. maculosus. In L. maculosus decipiens the flange (on its rare
occurrences) begins anteriorly at about the level of the mid-lateral
elytral macula, and ends well before the elytral apical truncation (fig.
278). Females of L. m. shermani, on the other hand, have the flange
beginning well behind the mid-lateral spot, and often it extends close
to the apical truncation (fig. 277). The condition for intergrades
appears to be intermediate.

Another modification of the female epipleura occurs in L. quadri-
lineatus, L. sonorensis, and L. vacaensis. In other species the median
margin of the epipleura is an unbroken curve even in those females
with flanges, but the females of these three species have an abrupt

JAMES R. ZIMMERMAN Lal

break in the epipleural margin at about the level of the first abdomi-
nal sternite. There may or may not be a flange present.

It is difficult to assess the phylogenetic importance of the modifica-
tions of the female epipleura in North American Laccophilus. Since
it is so obviously a highly variable sexual modification, it seems un-
likely that it can be used to show relationships among species or
species groups. Even races may differ significantly in the expression
or even presence of the character. It is evident, however, that the
modification in L. quadrilineatus, L. sonorensis, and L. vacaensis does
represent more specialization than do the differences in size or posi-
tion of the flange. One is inclined, therefore, to think that this char-
acter does support the view that those three species (and L. inagua
Young of the Bahamas) should be placed together in any grouping of
North American species.

11. Metasternum and postcoxal processes. There are some slight dif-
ferences in the curvature of the suture between the metasternum and
the metacoxal plate and in the shape of the postcoxal processes (fig. 2).
The plate for each species has an outline drawing of the metasternum
and the postcoxal processes. Comparable anatomical parts are all
drawn to the same scale. The most obvious difference that results
other than size can be seen in a comparison of L. quadrilineatus and
L. mistecus. Although the distance from the hind margin of the
mesocoxal cavities to the tips of the postcoxal processes is almost the
same in the two species, the distance from mesocoxal margin to the
posterior apex of the metasternum is greater in mistecus than in quad-
rilineatus (about 13 percent) — causing the internal lamina of the
metacoxae to be relatively shorter in the former. Other species simi-
larly can be compared.

There are also differences in the shape of the postcoxal processes.
One extreme is seen in L. maculosus (fig. 2) in which the processes
are strongly produced posterio-laterally, causing them to project well
posterior to the midline. The other extreme is seen in L. horni (fig.
209) in which the processes are not at all produced; and the midline,
or a point just off the midline, is the most posterior part.

12. Last visible abdominal sternite. Sharp (1882a, 1882b) frequently
made use of the last ventral segment in his descriptions of Laccophilus
species. It is generally a dimorphic character, but the degree of di-

MEM. AMER. ENT. SOC., 26

12 THE GENUS LACCOPHILUS

morphism varies from almost none in L. schwarzi or L. oscillator to
a large amount in L. ovatus or L. spergatus. Most males have an
asymmetrical curving crest on the sternite, while females have one
which symmetrically broadens and flattens anteriorly. Enough simi-
larity exists among species so that this segment can be used in only a
few diagnoses; but in such cases as the last two cited above, it can
serve for positive identification without examination of the aedeagus.
Sharp described three general types of ventral segments; a) entire
without sinuation on either side of the middle, b) with sinuation, and
c) truncated. In this paper type a would occur in L. youngi or L.
schwarzi (figs. 236-237), type b in L. pseudomexicanus males (fig.
84), and type c in L. spergatus (figs. 100-101). While these types
can be distinguished by description, it seems better to rely upon com-
parative drawings.

There is some phylogenetic significance in the expression of this
character, but interpretation is difficult. Too often there is too much
general similarity among species that are unrelated on other grounds.
One exception is the case of L. gentilis and L. ovatus, and another 1s
that of L. mistecus and L. spergatus. The last ventral sternite is
strongly modified in both instances and strengthens other anatomical
resemblances. L. gentilis and L. ovatus have spines along the lateral
sternal margins — a feature that occurs in none of the other North
American species. The spines, coupled with highly modified hind
margins, do indicate affinity of the species to one another and their
separation from the other species.

13. Legs. The first three segments of the male pro- and mesotarsi are
enlarged in a dorsoventral plane, and in a lateral plane as well as in
a few species. The degree of enlargement varies from easily seen
to barely detectable at 20 power magnification. The segments are
adorned with specialized hairs which are flattened distally into suck-
ing discs, or palettes. There is a row of four palettes on each of the
three segments. They function to help the male maintain purchase
during copulation.

Each species description gives the length of the fifth segment ex-
pressed as so many times the length of the fourth. The fifth tarsal
segment on the two front pair of legs is elongated in a few species —
more than twice as long as the fourth. An interesting correlation

JAMES R. ZIMMERMAN 13}

with this character is found in a modification of the female elytra.
The length of the male fifth segment is approximately equal to the
width of the anterior part of the female epipleuron. This means that,
when he grasps the female, his claws hook around the ventral edge of
the epipleuron and the tarsus bends at the seam between her epi-
pleuron and elytron or between his fourth and fifth tarsal segments.
His palettes then fit snugly against the dorsal surface of her elytron.
The male fifth segment of L. vacaensis has elongated until it exceeds
the width of the female epipleuron. To compensate, a crease (fig.
179) has developed above the epipleuron and now serves as the ful-
crum for the flexing of the tarsi. The crease is also weakly present in
a few other species.

Modification of the male tarsi is related to the nature of the micro-
reticulation of the female elytra. The tarsi are most enlarged in the
species in which the microreticulation is single and has a pebbled ap-
pearance (L. vacaensis and L. quadrilineatus). They are least modi-
fied in those species which have the microreticulation most clearly
doubled and have smooth and shining elytra (L. oscillator, L. schwarzi,
and L. duplex); and they are intermediate for others (L. maculosus,
L. proximus).

Another character of limited taxonomic value is the nature and
number of the spines or setae located on the proximal hind margin of
the pro- and mesofemurs (fig. 2). The mesofemoral setae are nor-
mally longer and thicker than the profemoral ones. The usual num-
ber of setae is about six on each femur, but in a few species it may be
as high as 12 on one or the other. The mesofemur usually has the
higher number. There does not appear to be any consistent pattern
to the exceptional numbers.

Attention was given to the hind legs, but I was generally unable
to find reliable characters that showed differences among the species.
An exception was L. youngi. This species has a greater difference in
length between the inner bifid metatibial spine and the outer one than
other species.

14. Male genitalia and oval plate. The structures include a) the
aedeagus, or intromittent organ; b) a pair of lateral parameres; and
c) a modified remnant of the ninth sternite, the oval plate (F. Bal-
four-Browne 1932). The aedeagus is the most useful diagnostic

MEM. AMER. ENT. SOC., 26

14 THE GENUS LACCOPHILUS

structure in Laccophilus. It is usually an elongate tapered organ
which bends about 30 to 60 degrees at near half its length, or curves
gradually over its whole length. It may also be twisted or have pro-
tuberances somewhere along its length. When two aedeagi are com-
pared side by side under the microscope, even slight differences in
conformation can be detected. On the other hand, the parameres and
oval plate, which may be equally sensitive diagnostic characters, are
more difficult to compare among species. They are described and
figured for each species, but they serve mainly to support conclusions
based on the aedeagus.

Species show little variation in the aedeagus through ranges that
cover hundreds or thousands of square miles. For example, males of
L. proximus from Florida, Indiana, and Nebraska have essentially
identical aedeagi even though some individuals may vary widely in
size, color, and elytral pattern. L. proximus presently has no races
recognized within it on the basis of differences in aedeagi or in other
morphological characters. Other species with well-defined races (such
as L. maculosus and L. pictus) have the same uniformity of structure
within races. The shape of the aedeagus is distinctive and consistent
in each race throughout the range, except for intermediates that occur
in intergrade zones. Intergradation of the aedeagus adds more sup-
port to the weight given it in taxonomic determinations.

Certainly, one can find regional differences in aedeagi if enough
time is spent and enough comparisons are made, but without discon-
tinuity in structure no taxonomic distinction is necessary. Consider
L. mexicanus with its three races. The only reliable difference be-
tween L. m. mexicanus and L. m. atristernalis is the slightly different
tips of the aedeagi. As yet, their known ranges are narrowly allo-
patric in southern California with no intergraduation demonstrated;
but, within the range of each, the aedeagus remains the same. Even
slight differences in aedeagi can be indicative of sufficient change in
genotype to warrant taxonomic recognition. The third race, L. m.
oaxacensis, differs in several characters besides the aedeagus. It ap-
pears to be completely allopatric from L. m. mexicanus and could
have been described a different species instead of just a race. I have
preferred to retain it within L. mexicanus. The same reasoning ap-
plies for L. gentilis, L. quadrilineatus, and L. peregrinus.

It does not follow, however, that species separation will not result

JAMES R. ZIMMERMAN iI)

without differences in aedeagi since reproductive isolation is the basis
for determination of species limits and morphological differences
merely serve to detect those limits. I am unable to distinguish be-
tween the aedeagi of L. quadrilineatus and L. sonorensis, and they
normally would be considered a single species; but there is a qualita-
tive difference between elytral patterns with no intermediates in a wide
area of sympatry. Here other characters have precedence, and the
populations are considered separate species. Polymorphism may be
involved instead of reproductive isolation, but that has not been tested.

As yet I have not been able to see broad phylogenetic significance
in comparisons of male genitalia.

Quantitative characters. — Four size measurements were used for sta-
tistical comparisons of populations. They are |) total length, 2) ely-
tral length, 3) total width, and 4) pronotal width. All can be taken
from a dorsal view through an American Optical Cycloptic stereo-
scope fitted with either a 400 square grid or a reticle divided into 100
increments. Both were used on occasion. One grid square was equal
to five increments and one increment was 0.054 mm at 20 power
magnification. Measurements could be reliably interpolated to 0.5
increments.

Total length is the distance from the front of the head to the tip
of the elytra. It does not include the tip of the abdomen which may
or may not protrude. The measurement can be replicated if both
endpoints are sharply in focus. Parallax error is reduced if the speci-
men is positioned horizontally instead of vertically in the field. Parted
elytra, distended neck, and upturned head may contribute to inaccu-
racy. Normally the elytra should be joined, the head be drawn tightly
into the pronotum, and the mouthparts facing ventrally for uniformity.

Elytral length is the distance along the elytral suture from the
apex of the pronotum to the elytral tips. Parted elytra and parallax
errors contribute most to mismeasurement. The elytral surface curves
upward, causing trouble sometimes in distinguishing the exact inter-
section of the elytral suture and pronotal apex. The specimen must
have the head and elytral tips on the same plane.

Total width is the greatest distance across the body and occurs
just behind the midpoint. Frequent separation of the elytra gives this
measurement limited use. The sample number is often less for many

MEM. AMER. ENT. SOC., 26

16 THE GENUS LACCOPHILUS

localities, and the measurement is not included in a few cases. The
value is more reliable for females than males, because the latter were
manipulated more in curation. Teneral (recently emerged) specimens
were omitted from the sample because of their shrunken condition.

The most reliable of the four quantitative characters is pronotal
width. It is the greatest width across the base of the pronotum. It is
little affected by distortion or disturbance of other body parts. Its
endpoints are sharply defined and can be simultaneously adjusted into
sharp focus.

A fifth character is also included in the tables. It is a ratio ob-
tained by dividing pronotal width by elytral length. It is frequently
a remarkably consistent character for different populations. Values
are virtually the same to three decimal places.

Each species has a table which contains sample size, arithmetic
mean, and standard deviation for all measurements from different lo-
calities. Also included are the rational means with their 95 percent
confidence intervals. The locality should be interpreted as the center
of the area from which the specimens were taken. The areas of geo-
graphical localities were restricted as much as possible, but some
lumping from several localities appeared advisable in order to obtain
larger samples. Decisions were necessarily arbitrary in those cases.
I have not described any species or races merely from quantitative
characters. More refined sampling and more experience in selecting
characters is required for individual species problems.

AFFINITIES AND DISTRIBUTION PATTERNS
OF NoRTH AMERICAN LACCOPHILUS

There are five or six separate phylogenetic groups within the 27
species of North American Laccophilus. Four of them, comprising
19 species, are relatively easily characterized; i.e. their members share
a complex of characters and, at the same time, can be separated from
other species. The other eight present a more confused situation, and
it is not possible as yet to reliably draw boundaries for inclusion and
exclusion.

The species groups on the North American continent appear to
have several origins. The maculosus group seems to have originated
and developed on this continent; L. biguttatus clearly has come from
the Palearctic Region; the gentilis group are invaders from South

JAMES R. ZIMMERMAN iL7/

America; and the vacaensis group are, while tropical, not clearly of
South or North American origin. At the same time L. undatus and
L. schwarzi appear to be relict species of the temperate Northeastern
deciduous forests, having close relatives only much farther to the south.

The maculosus group includes the largest number of species —
nine, which are L. maculosus, L. fasciatus, L. proximus, L. salvini,
L. mexicanus, L. spergatus, L. mistecus, L. pseudomexicanus, and L.
fuscipennis. Most of them contain several well-differentiated races.
The species cover wide geographical areas and are often extremely
abundant. They are perhaps the most successful dytiscids in North
America. All are irrorated species, but some may also have irregular
black markings on the elytra. Five of the species have black on the
venter, and the other four have venters that range from brownish-
yellow to reddish-brown. Males have metacoxal files (some females
also have traces), and the females have a fine saw-toothed ovipositor.
Other characters are the strong dorsoventral enlargement of the male
pro- and mesotarsi, the produced postcoxal lobes, and an intermediate
condition of the elytral microreticulation; i.e., tending toward double.
Although the species are exceedingly similar, male genitalia provide
excellent characters for discrimination. Females of some species are
so much alike, however, that considerable experience is necessary to
distinguish among them; and, in some instances not even experience
allows complete assurance.

The distribution of the maculosus group suggests an origin in
North America. The largest concentration of species occurs on the
Plateau and Central Highlands of Mexico. Eight of the nine species
have been taken in this area. L. fuscipennis and L. proximus more
typically are restricted to lowlands, but in the United States, L. proxi-
mus does commonly occur on the Great Plains at up to about five
thousand feet. Routinely, four or five species of the maculosus group
occur together in central Mexico. From this high concentration the
number of sympatric species declines in a roughly concentric pattern
to one species in the state of Washington, two in southern Baja Cali-
fornia, two in Florida, two or three on the Yucatan Peninsula, and
two in South America.

One might reason that there is more available habitat in the area
of high frequency, but the large number of other Laccophilus species
outside of highland central Mexico argues against that conclusion.

MEM. AMER. ENT. SOC., 26

18 THE GENUS LACCOPHILUS

The high frequency seems to be more easily explained by assuming
that central Mexico is the center of origin for the group and that they
have spread out from that area.

Two other species with metacoxal files show some affinity for the
maculosus group; but neither are irrorated and both have a more
strongly double microreticulation and, consequently, are more shining.
L. pictus is brightly colored with a strongly contrasting yellow and
black pattern. Those characters, along with the differently shaped
pronotum, argue against placing it in the maculosus group. The range
of L. pictus is completely contained in North and Central America,
however; and it might be a radically changed offshoot of the macu-
losus group. The second species, L. youngi, differs from the maculosus
group only in its marmorated rather than irrorated pattern and micro-
reticulation. It is clearly a Neotropical species and may provide a
link between the maculosus group and species from the Andean
highlands.

The next largest species group is the one which includes L. vaca-
ensis and L. quadrilineatus. Other species included within this group
are L. spangleri, L. peregrinus, L. huastecus, L. raitti, and L. sonor-
ensis. The recognition of the existence in North America of this
assemblage is one of the more surprising developments of this study.
Formerly only vacaensis and quadrilineatus were known, and those
were known only as monotypic species; but, in reality, the group is
composed of at least seven species. Four of these contain two or
three subspecies.

Two characters, both in the female, indicate the affinity of the
members of this group. The first and most striking is the large toothed,
rakelike ovipositor; and the second is the single condition of the ely-
tral microreticulation which causes the females to be less shining and
to have a somewhat velvety appearance. Other characters are the
lack of a metacoxal file in the male and the strongly modified male
pro- and mesotarsal segments. These segments are strikingly enlarged
in a dorsoventral plane; but, in three species, there is also lateral ex-
pansion. The male aedeagi show less modification among the species
than is seen in the maculosus group. Elytral patterns vary among
irrorated and marmorated and some intermediate condition. L. hu-
astecus, L. peregrinus, L. raitti, and L. vacaensis can be either and
vary both racially and individually. L. quadrilineatus and L. sonoren-

JAMES R. ZIMMERMAN 19

sis are marmorated; and L. spangleri is irrorated, but with a strong
tendency for fusion of the dots. Some individuals of L. v. vacaensis
are superficially very close in their elytral pattern to proximus of the
maculosus group (they are sympatric), but the males are easily sepa-
rated between the two species.

The third group is of South American origin and contains but two
species in North America. It is the gentilis group which contains L.
gentilis and L. ovatus. L. gentilis has at least two races — one in the
extreme southeastern United States and the other in all tropical low-
land regions from Mexico to Panama and presumably beyond. It is
also in the West Indies as L. bifasciatus Chevrolat and perhaps as
some other species with which I am not acquainted. L. ovatus, which
was described from South America, is racially different from its North
American representative. The close relationship of L. gentilis and L.
ovatus can be seen in several characters which, at the same time, show
them to be distinct from other North American species. The most
important is probably the long, acuminate prosternal process. Only
these two species among the North American species have this con-
dition. Another unique character is the presence of an irregular row
of short spines or setae along the lateral margins of the last ventral
abdominal segment. Both species tend to be more parallel-sided than
other Laccophilus. In addition, both have shining elytral surfaces
(microreticulation clearly double) which are generally brownish-red
above with undulating yellow, subbasal bands and usually more yel-
low subapically. The genitalia are highly distinctive in contrast to the
strong superficial resemblance of the two species. The ovipositor has
fine sawteeth.

The presence or absence of a metacoxal file is not particularly im-
portant in this group. A file is present in one race of L. gentilis, but
is absent in the other and in L. ovatus. This situation could readily
be viewed as reinforcement or character displacement. In Mexico
and Central America L. ovatus is sympatric with L. gentilis suavis,
and the file is well-developed in the latter. But in Florida L. g. gen-
tilis retains only a trace of it. L. proximus and L. fasciatus, filed
members of the maculosus group, are the common species of Lac-
cophilus in Florida. They also occur in Mexico with gentilis but not
commonly in the same habitats.

MEM. AMER. ENT. SOC., 26

20 THE GENUS LACCOPHILUS

The gentilis group represents a small North American extension
of what appears to be a much larger group in South America which
probably includes L. latifrons Sharp, L. planodes Guignot, and L.
curvifasciatus Guignot among others.

L. biguttatus appears to be the single Palearctic representative that
has successfully migrated to North America. It closely resembles
L. minutus Linnaeus and could easily be considered a race of that
species. L. biguttatus is the only boreal species of Laccophilus, a fea-
ture that agrees with the distribution of L. minutus in Eurasia. It has
been able to extend its range into California via the Sierra Nevadas
and into northwestern New Mexico via the Rocky Mountains. In
spite of a wide distribution from coast to coast in northern North
America, L. biguttatus has not undergone any marked racial differ-
entiation, which supports the view that it is a recent immigrant.

The last six species of North American Laccophilus form a hetero-
genous assemblage which cannot easily be assigned to a single phylo-
genetic lineage or to one of the previous groupings. They do have
characteristics which show some affinities among themselves. None
are irrorated, but instead have pigment suffused over the elytra form-
ing yellow spots or irregular, often interrupted bands surrounded by
black, reddish brown, or some admixture of those colors. Metacoxal.
files are lacking, all have fine, sawtoothed ovipositors, the elytral sur-
faces are shining with strongly doubled microreticulation, and the
prosternal process is either short or intermediate.

They separate into two distinct species pairs and two problemati-
cal species. The first contains L. horni and L. leechi which resemble
one another so closely that one is inclined to consider them as races of
a single species; but it is difficult to decide whether they are allopatric
or sympatric, and they have accordingly been retained as separate
species. Their most outstanding features are the brightly spotted yel-
low and black elytral patterns, the prominent pigmentation on the
pronotum, and the truncated postcoxal processes. They show no
particular relationship to any one of the other four species, however.
Their range is largely confined to Mexico north of the Isthmus of Te-
huantepec, and to the southwestern United States. L. leechi is the
more restricted of the two and has been found only in two locations in
the lower mountains on the West Coast of Mexico.

L. oscillator and L. duplex are another species pair that are very

JAMES R. ZIMMERMAN Dil

similar to one another, but sympatry has been established. The ely-
tral pattern is the suffused type with an irregular undulating subbasal
band and an incomplete postmedian one. L. oscillator has a short
prosternal process; and L. duplex has an intermediate one, but not
nearly as long as that of L. gentilis. L. schwarzi also has an inter-
mediate length prosternal process, the same general type of elytral
pattern, and an aedeagus similar to L. oscillator and L. duplex so that
it may be related to those two species. They are tropical species which
extend only as far north as Nuevo Leon and Tamaulipas (L. duplex)
and Arizona (L. oscillator). Both apparently extend into South Amer-
ica and the West Indies. L. schwarzi has been collected only in the
vicinity of Washington, D. C., and Amherst County, Virginia. It
could be a relict of a population that formerly extended much farther
south. L. schwarzi may in turn be close to L. undatus, but no charac-
ters place it closer to L. undatus than to L. duplex (in fact, the reverse
is true). LL. undatus is confined to the northeastern hardwood decidu-
ous forest belt. It is a unique species that cannot be closely allied
with any other North American Laccophilus and surely must be a
relict. It would not be too surprising to find it has a close relative in
the temperate Andean highlands.

L. undatus, L. schwarzi, L. duplex, L. oscillator, L. horni, and
L. leechi do share one common ecological relationship that separates
them from other North American Laccophilus. Instead of preferring
small aquatic habitats in sun-exposed environments, they all prefer
shaded habitats and, more often than not, pools or streams rather than
ponds or ditches. This could represent the beginning of a gradistic
change that may be more significant than any presently detectable
morphological difference that sets them off from other Laccophilus.

Summary. — Construction of a phylogenetic tree of North American
Laccophilus would be premature at this time. It will be necessary to
have a broader view of the total diversity within the genus and more
reliable criteria for which are primitive or derived characters within
the genus. Only one character seems to be a fairly good indicator of
specialization in the North American species — the wide-toothed rake-
like ovipositor of the vacaensis group. Other shared characters seem
to be just as attributable to parallelism or convergence as to affinity.
It will probably require studies of internal anatomy, cytology, and

MEM. AMER. ENT. SOC., 26

i)
tN

THE GENUS LACCOPHILUS

immature stages —as well as external morphology — to arrive at reli-
able criteria for primitive and advanced characters.

Nearly all members of Laccophilus appear superficially similar
due mainly to their streamlined shape; but there can be no doubt that
there has been considerable divergence in them, as the North Ameri-
can species demonstrate. They are a highly successful group, and
some species are perhaps expanding their ranges and also undergoing
a vigorous speciation. The species of North America derive from
several sources: 1) The Old World (L. biguttatus), 2) South Amer-
ica (the gentilis and oscillator groups), and 3) North America (the
maculosus group). The vacaensis group are morphologically the most
homogenous and could be considered as subgenerically differentiated,
but this should not be done until the South American species are
better known. The horni-oscillator group, although not morphologi-
cally distinctive or even homogeneous, have diverged ecologically to-
ward a different type of aquatic habitat.

Most Laccophilus species are wide-ranging, outbreeding forms
which can survive in a variety of aquatic habitats; and an under-
standing of North American Laccophilus is only possible after the
South American and West Indian species are better known.

Genus LACCOPHILUS Leach

Laccophilus Leach, 1817, p. 69; Type species: Dytiscus minutus Linnaeus, 1758
(= Dytiscus obscurus Panzer, 1794); fixed by Westwood, 1838.

DIAGNOSIS. — The North American members of this genus are small
(3.0 mm to 7.0 mm), streamlined, shining insects that are most easily recog-
nized among the dytiscids by the produced lobes on the hind margins of the
metatarsal segments (fig. 1). The closely related, but much rarer genus, Lac-
codytes, has produced lobes also, but it differs in having simple, instead of
bifid, metatibial spines, and the largest Laccodytes probably never exceeds 2.5
mm. The head of Laccophilus appears extremely short when viewed from
above due to the generally hypognathous attitude. The head, pronotum, and
elytra form a continuous outline; and the tip of the prosternal process is uni-
spinose. The males of about a third of the species have a series of short ridges
(metacoxal file) on the metacoxal plates. An excellent field character is the
manner in which they jump when captured in a sieve. They are able to hop
a distance of one or two inches when attempting to right themselves. Large
dytiscids, such as Dytiscus and Cybister, will do the same thing; but few, if any,
of the small ones do.

DESCRIPTION. — General appearance: small (3.0 to 7.0 mm), oval, shin-

JAMES R. ZIMMERMAN 23

ing; head, pronotum, and elytra with continuous streamlined outline; widest at
about or just behind the middle; colors ranging from black and brown to red
and yellow and mixtures thereof. Head: hypognathous, extremely short in
dorsal view; incomplete line of punctures just above the clypeal margin. Pro-
notum: posteriolateral angle rectangular or obtuse; not margined. Elytra: scutel-
lum hidden; elytra varying from slightly truncated to slightly attenuated; some
females with a flange flaring laterally from the posterior part of the epipleura;
posteriolateral margins fringed with long posteriorly directed hairs. Micro-
reticulation: fine polygonal mesh over dorsal surface; secondary mesh some-
times formed due to weakening of smaller inner cellules; coarser in females;
cellules frequently elongate, especially on venter. Venter: prosternal process
with ventral crest, widened in apical third posterior to procoxae; expanded part
varies from diamond-shaped to strongly acuminate; episternum of metathorax
does not reach the mesocoxal cavity; metasternal wings very slender, arcuate;
males of some species with a metacoxal file —a series of ridges curving pos-
teriorly and laterally away from the midline; postcoxal processes varying from
slightly rounded and produced to truncated; last ventral abdominal segment
may be rounded, truncated, or notched; oblique striations on all abdominal seg-
ments; setigerous punctures scattered over surface of last segment, usually thick-
ened toward hind margin. Legs: first three segments of pro- and mesotarsi
of males enlarged in at least a dorsal-ventral plane and fitted with a series of
rows of sucking discs; claws simple and equal on front two pair of legs, unequal
on metatarsi; antennal comb on ventral margin of profemur; conspicuous setae
on basal ventral margin of pro- and mesofemora; metatarsi with produced lobes
on hind margins; metatibial spurs bifid, unequal; two or three large spines lo-
cated on metatarsal posterior margins. Genitalia: aedeagus sclerotized, at least
twice as long as either paramere; right paramere more strongly sclerotized than
left, roughly triangular, and with one or two hairs projecting from near apex;
left paramere usually larger than right, heavily sclerotized at base, but more
flexible and fiattened apically; apex rounded with one or two hairs; oval plate
with median ventral ridge and usually an acuminate tip; ovipositor toothed
ventrally, strongly sclerotized.

CHECKLIST OF THE LACCOPHILUS OF NORTH AMERICA

1. maculosus

m. maculosus Say (= maculosus Germar) new synonymy and status
m. decipiens LeConte new status
m. shermani Leech new status

2. fasciatus

f. fasciatus Aubé (= apicalis Sharp) new synonymy and status
f. rufus Melsheimer (= fasciatus Aubé) restored name, new status
f. terminalis Sharp new status

MEM. AMER. ENT. SOC., 26

Ww

10.
11.

13%
14.

15.
16.
fs

THE GENUS LACCOPHILUS

proximus

p. proximus Say
p. americanus Aubé

salvini Sharp

mexicanus

m. mexicanus Aubé
m. atristernalis Crotch
m. Oaxacensis

pseudomexicanus

mistecus

m. mistecus Sharp
m. aztecus

spergatus Sharp
fuscipennis Sharp
youngi
pictus

p. pictus Castelnau

p. coccinelloides Régimbart

p. insignis Sharp
vacaensis

v. vacaensis Young
v. chihuahuae
v. thermophilus

spangleri
peregrinus

p. peregrinus
p. variabilis

huastecus

raitti

quadrilineatus

q. quadrilineatus Horn
q. tehuanensis
q. mayae

sonorensis
biguttatus Kirby
horni Van den Branden

leechi

new
new

new
new
new

new

new
new

new

status
status

status
status
subspecies

species

status
subspecies

species

status
status
status

status
subspecies
subspecies

species
species

subspecies
subspecies

species

species

status
subspecies
subspecies

/ species

species

Dike

JAMES R. ZIMMERMAN WS

undatus Aubé
schwarzi Fall
duplex Sharp (= optatus Sharp) new synonymy
oscillator
o. oscillator Sharp new status
o. laevipennis Sharp new status
gentilis
g. gentilis LeConte new status
g. suavis Sharp (= championi Sharp) new synonymy and status
ovatus zapotecus new subspecies

KEY TO THE LACCOPHILUS OF NORTH AMERICA NORTH OF PANAMA

NM

AND EXCLUSIVE OF THE WEST INDIES

Elytra with irrorations (color applied as grains of sand) ................00 2
1 EVVAURE, WAIHI OYOYUE TTAROTENSTOYOS” ccscosccs0cb0es200000000000005000H0600e50050000000d0007 000380020500 14
Males with metacoxal file (fig. 2); females with sawlike ovipositor (fig.

(AND) Re sen asc ScooS RESO Eee SEOFEE COSC Cr ECE ACE EOD ECCOOREECOE Roce er Ree EO Ree Coo 3
Males without file; females with rakelike ovipositor (fig. 188) .......... 10
Venter with black or nearly black (best viewed megascopically) ............ 4
Venter with various shades of yellow, red, or brown ................::::eeeees 5

Elytra evenly irrorated; last ventral segment of males strongly truncated
(fig. 100); that of females slightly truncated (caution: do not confuse
translucent spots as concentrations of COlOr) .................ccceeeee spergatus

Elytra unevenly irrorated (see table below). All of these and spergatus
are easily separated from one another by the male aedeagus but are
superficially extremely similar ...............c:.ccccsssceceesseees salvini, mexicanus,

pseudomexicanus, mistecus

Elytra tending to form an irregular black fascia across its posterior half
(extends anteriorly also in f. fasciatus), sometimes appearing checkered

wheniviewedmmesascopicallivamessssssseceseerectes ee erere entrees fasciatus
Elyira with OU ta tasclaynsscecccccessessacc es cea teres Geof OS Sea CE Ts wees hes ba eR Sabato 6
Three or four large, well-defined maculae along the lateral margins of the

CIV IRE ccoseenquoacecece3acpeccdcGoosac SoScGBee Ho UCODEERE ce SBE CocoE SSG sc EESCo SEE maculosus
Pateralemacnlacenotawell-delime dines. cecssseresce eran ee ceeneeee ee nee eee enersemcens 7
rv Crea Gna sae cece cee ene et Reco Coe ae dee ecate shae ceaurehsWeadascevttne ceases 8
COKWEIe OTRO canaetanscseeccaccoqenpacecod eas cts eau Ree cee eee EERE mistecus aztecus
Coastal regions of California and Oregon ................ mexicanus atristernalis
INOtroccuming ny Califommiavand Ore gonyercscccserescresece sec esereneee eens 9

Elytral pattern with limits easily distinguished in anterior half; males with
strongly developed file and female with weak one; broadly distributed east
of Rocky Mountains and east coastal lowlands of Mexico ...... proximus

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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JAMES R. ZIMMERMAN Di

Pattern limits not easily determined; file weak in male: Mexican lowlands

BSE SAECO CaGEOO COOGEE COG ICBO UOC EECHOCRE OTE DECREE EEE UEC EE Cee ae Ree CPeeer fuscipennis
Front of head darkened; irroration indistinct at best .................... huastecus
Occiput, but not front of head sometimes darkened; irroration strong or

IW Geek Kate sccr cera vaccaceeecetee cuce cee ces ae cec ce tvusen odes eeuseul eens decstvnceoUessusvescuceactaesBectees 11
Irroration indistinct at best; complete suffusion of color in many speci-

MNLCIUS as cececetccvcsreesssessay sn saaetsct ces todece cuss sileivy sess sevecksconsueudvassescecdeesssevesases il72
Irroration distinct on most of elytral pattern .0...........ccccccceeeeseeeeeeeeeeees 13
Well-defined crest above epipleural seam in females (fig. 179); length 4.8

LOPEZ MIMI SATe bs AS smINAV ATI ty ececccceccecceseccses oreeresseste-cossesscccateocceceees raitti
Poorly defined crest above epipleural seam in females (fig. 172); length

3.8 to 4.3 mm; Neotropical Mexico .............ceeeeeeee peregrinus peregrinus

Male with fifth pro- and mesotarsal segments two and one-half to three
times as long as fourth; female epipleuron abruptly narrowing just be-
hind level of hind coxal cavities and frequently with flange (fig. 277);
length 4.3 to 4.9 mm; southern United States to Panama and West
10 Tei reer tcccrecnenccstcetarccoctectad eee eer ee ceeee Mee ce re ere corre reerre vacaensis

Male fifth pro- and mesotarsal segments no more than twice as long as
fourth; female epipleura narrowing gradually (fig. 2); rarely, if ever
with flange; length 3.9 to 4.8 mm; Veracruz to Nicaragua ...... spangleri

Elytra uniformly brownish-yellow or light brown; apical segments of an-
tennae and palpi darkened; length 3.9 to 4.8 mm; Canada, and high
mountain areas of the northern United States ................00.000000 biguttatus

Not as above (antennae and palpi of undatus darkened) ..............0000+ 15

Prosternal process long; i.e., groove on mesosternum which receives process
extends well posterior to mesocoxal cavities (fig. 261); lateral margins
of last ventral segment with irregular row of short spines or setae .... 16

Prosternal process short; no spines present on lateral margins of last ven-
tale SE LIME tagcrececcceceseesee esos sees e ree cease. Rasen Ae esetneesderssTalaevsadsosehovee 17

Male without metacoxal file if occurring in the United States, but with file
in Latin America; apex of last ventral segment not produced; reddish-
brown elytral color evenly applied; length 3.0 to 3.5 mm; Gulf Coast,
DONO AM OME AM ANN alee neces evens cee: soca cwekae cee ca ences Seneca edie e Bec bul ees utehe gentilis

File absent; last ventral segment truncated and with produced apex (figs.
270-271): elytra uneven and frothy; reddish-brown color; length 3.3 to

3.8 mm; Veracruz and Yucatan to South America .............:.00e ovatus
Elytra strikingly marked with yellow spots on zigzag stripes surrounded by
EYE. Gir COCOA IDO osc5cc000009300005060070000003500400000260000089050000055 500050000006 18
IN OUPASTADON EN cesscces ster seassecnesaticese seer ccscaseeesstcseviccestodterrsosetecsosecacsesssaesosersse 20

Postcoxal processes laterally produced (fig. 2): male with file; metacoxal
plates entirely pale; epipleura strongly darkened in anterior half; length
4.5 to 6.0 mm: southwestern United States, Baja California, Mexico to
(COS TAMRAC A eee eer ce Ra sone Meck neuen sty: duscabonrav seers semesebevae ee es pictus

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

Postcoxal processes truncated (fig. 209); no file in males; metacoxal plates
darkened at least on lateral margins; epipleura weakly darkened at most

REE Cera CoCo beac doen dobes oscar erin nccann todosaseacdsoobnoctastpbncaceceodon0006000 19
Dark pronotal markings on front and base confluent across disc; length 3.5
to 3-8 mm) Nayanityand ialiscomcreresseseescareeerterteceeerer ree leechi

Pronotal markings not confluent across disc; basal one suggestive of handle-
bar moustache; length 4.2 to 4.8 mm; Arizona and Texas to Oaxaca ....

Lie Sal da abe De gend caau un dovetle eneet eur usee Pan cat ate cereece rel receanon ULCiH Ia nar ae o horni
Occurring in eastern United States ...........ccccsssesessseeecceceeeeceeeenseenseeees 21
Not extending east of Texas and north of Kansas in United States ...... DD,

Tips of maxillary palpi and antennae darkened; brown, yellow, and black
variegated elytral pattern; lateral margin of metacoxal plates not dark-
ened; length 3.2 to 4.3 mm; northeastern United States ............ undatus

Palpi and antennae not darkened; lateral margin of metacoxal plate dark-
ened; elytra brownish-red with incomplete subbasal band; Washington,

Dis and! Virginia oo cccystussseassedecees casecae s eeseeds on eee eater Ree schwarzi
Front of head darkened; length 3.9 to 4.2 mm; Veracruz ............ huastecus
Erontof head notidarkemed i yesccesessee cesar ee 23

Elytra with irregular subbasal band; surface highly shining; postcoxal proc-
esses truncated (fig. 209). (Note: male genitalia must be examined to
satisfactorily separate following speCieS) .............::ccccceeqeessteeeeceeeeneneees 24

Elytra marmorated or variegated, but without subbasal bands; surface micro-
reticulation single; postcoxal processes laterally produced (fig. 2) ...... Ds)

Darkening of pronotum extending from base to disc; sternites usually dark-
ened; length 4.1 to 5.8 mm; Arizona to Panama ...............:::005 oscillator

Darkening confined to base and not attaining discal region of pronotum;
sternites pale; length 3.5 to 4.1 mm; Tamaulipas to Costa Rica .... duplex

Male with metacoxal file; female with sawlike ovipositor (fig. 42); length

4.6 to 5.4 mm; Nayarit to Costa Rica .............ccccceceeeeeseeseesessetenees youngi
No metacoxal file; rakelike ovipositor (fig. 188) ..........cccesceecssseceenreeeees 26
Pro- and mesofemoral setae (fig. 2) at least twice as long as femur is wide

(species usually exceeding 5.0 MM) ..........cccccceccecccceeeeeeeeeeeeessessesnsenees 27
Pro- and mesofemoral setae less than twice as long as femur is wide (spe-

IES WS UENIK ESS neta SA0) se0t00\)) ” Goneooosedoooocodsopocosbadodconoc0esdco0500500000000000000 28

Marmorated elytral pattern covering most of elytra (Yucatan) or with
four relatively solid colored prominent anterior extensions from a post-
median blotch; length 5.0 to 6.7 mm; Kansas to California south to
Isthmus of Tehuantepec and Yucatan Peninsula .............. quadrilineatus

Elytra with four open anterior extensions from a postmedian blotch (fig.
306); length 5.0 to 6.0 mm; southwestern New Mexico to south Cali-

fornia, Sonora and Baja California ....................cccc0eeeseeeeeneeeeees sonorensis
Length 4.8 to 5.2 mm; females with well-defined crest above epipleural
seam (fig. 179); known only from Nayarit ............c::ccsssccssseceereeeeees raitti

Length 3.8 to 4.6 mm; poorly defined crest above epipleural seam in fe-
males (fig. 172); usually occurring south of Isthmus of Tehuantepec
ngesedersaoeSeoies suc chscadvs contest coseve ouscauestuedsssavescset seers teascecmeeeeeeess peregrinus (part)

JAMES R. ZIMMERMAN 29

LACCOPHILUS MACULOSUS

The three well-defined races in this widespread polytypic species
were previously considered species. Laccophilus was described by
Say in 1823 from the eastern United States (presumably from Indi-
ana); L. decipiens by LeConte in 1852 from California; and L. sher-
mani by Leech in 1944 from Arizona and Texas. Their present
ranges and intergrade zones suggest that they were geographical iso-
lates during the last glaciation. After the ice sheet retreated, second-
ary contact occurred in the northern Great Plains between maculosus
and decipiens. A similar contact between decipiens and shermani
took place in northern Arizona and New Mexico as the forests’ lower
margins retreated to higher elevations. Today maculosus is most con-
centrated and abundant in the glaciated region of the northeastern
deciduous forest. It is replaced by decipiens in the western United
States, southern Canada, and Alaska. This race in turn intergrades
in a narrow band with shermani, the southwestern and Mexican
replacement.

L.m. maculosus is smaller and more strikingly patterned than the
other two races, which superficially are extremely similar. It also has
less size difference between the sexes than is found in decipiens and
shermani. However, shermani shows divergence in secondary sexual
characters (metacoxal file, epipleural flange, and aedeagus) from de-
cipiens and maculosus which have diverged relatively little in those
characters.

Apparently, there is sufficient ecological separation so that inter-
grades can be successful only in narrow, intermediate zones. There
is no evidence of swamping. The intergrade populations manifest
enough heterogeneity in all characters to suggest some backcrossing
and F»2’s.

DESCRIPTION. — Large (length 5.0 to 6.4 mm; width 2.3 to 3.3 mm),
brown, irrorated species; metacoxal file present, prosternal process short, ovi-
positor sawlike. COLOR. Head: pale brownish-yellow above and beneath;
faintly darkened with reddish-brown at the base of the pronotum; appendages
pale yellow except mandibles that darken to brownish-red toward tip. Prono-
tum: pale brownish-yellow with reddish tint. Elytra: dark brown irrorations
on a pale brownish-yellow background; prominent pale spots present (figs. 1,
281); irrorations often in strings and tending to coalesce around maculae; epi-
pleura pale yellow anteriorly and sometimes with reddish-brown posteriorly.

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

30

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MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS
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JAMES R. ZIMMERMAN 33

Tergite VIII: pale yellow or light brown. Venter: prosternum, its process, and
hind legs pale yellow to reddish-brown. Genitalia: aedeagus, right paramere,
oval plate, ovipositor all reddish-brown; left paramere with yellow on distal
half. ANATOMY. Méicroreticulation: weakly double on head, pronotum, and
elytra; secondary mesh discernible, but individual cellules still apparent. Head:
supraclypeal seam closely parallel to margin. Pronotum: WH/PW, 0.66 to
0.68; LP/PW, 0.42 to 0.43. Elytra: female epipleura sometimes with flange;
rarely in decipiens and maculosus, but almost invariably in shermani (figs. 276-
281); truncation of apices slight. Venter: coxal file composed of about 25 to
35 lines; prosternal process with well-defined crest; lobes of postcoxal processes
rounded and laterally projecting well posterior to midline (figs. 2, 28); last visi-
ble ventral abdominal segment rounded and similar in males and females; that
of males slightly produced and with asymmetrical curving ridge in middle (fig.
30); last segment of females weakly arched with posterior margin not notice-
ably produced. Legs: male pro- and mesotarsi enlarged in a dorsoventral
plane; palettes easily distinguished at 20 power magnification; male fifth tarsal
segment on both pair of legs about one and three-quarters as long as corre-
sponding fourth; profemoral setae (about 6) shorter and finer than mesofemoral
ones (8 or 9). Genitalia: oval plate (fig. 32) large and produced to acuminate
tip; its ventral crest prominent and extending forward with little lateral curva-
ture about one-half the length of the plate; an indefinite number of raised lines
lying on either side of crest; right paramere about half as long as left, roughly
triangular in outline and heavily sclerotized throughout (figs. 33, 34, 38), with
one or two hairs projecting subapically; left paramere while sclerotized at base
is less so and more plate-like distally; also with one or two hairs projecting (fig.
38); ovipositor with about 13 to 15 sawlike teeth on each ridge.

Laccophilus maculosus maculosus Say, new status
(Biostelhn22.8-335385 2oL)

Laccophilus maculosus Say, 1823, pp. 100-101 (taken from J. L. LeConte’s
editing of the complete writings of Thomas Say, 1891, p. 514). Neo-
type: male, Bloomington, Monroe County, Indiana, Bloomington, 5 m. N.,
iv.26.56, J. R. Zimmerman, USNM; Blatchley, 1910, p. 209; Zimmerman,
1960, pp. 142-143.

Laccophilus maculosus (Germar), 1824, p. 30; Aubé, 1838, p. 42: Crotch,
1873, p. 399; Sharp, 1332a, p. 289; Wickham, 1895, p. 72; Leng, 1920,
p-. 76; Zimmermann, 1920, p. 21; Zimmerman and Severin, 1957, p. 29;
Zimmerman, 1960, p. 143; Gordon and Post, 1965, p. 12.

DIAGNOSIS. — Size and elytral pattern serve to separate m. maculosus
from related subspecies and other sympatric species in northeastern North
America. L.m. maculosus is the largest Laccophilus in the northeastern United
States and Canada. The unevenly irrorate pattern with strongly contrasting
maculae is a distinctive trait as witnessed by the same epithet given in the syn-

MEM. AMER. ENT. SOC.. 26

34 THE GENUS LACCOPHILUS

> Tergite VIll

Figure. 1. Dorsal view of Laccophilus m. maculosus male.

onymy. The pigment is concentrated around pattern margins and in areas
scattered throughout the disc. The accumulation of pigment immediately be-
hind the midlateral spots is suggestive of L. fasciatus rufus which has a post-
median stripe or blotch, but maculosus is larger (length, about 5.0 to 5.8 mm
as compared to 4.5 to 5.1 mm) and the anterior irroration is more marked.

JAMES R. ZIMMERMAN 35

antennal comb Cre es
fie
bg Oe

palettes

mesosternal epimeron
metasternal episternum
prosternal process

mesocoxa
“pmetasternal wing

profemoral setae

-tepipleuron
mesofemoral setae ;

4 metacoxal file

¥
Hist visible

g Oa abdominal segment

Ht postcoxal process

Wy, HA

metatibial spurs

6th visible

metatarsal lobes _ ae abdominal segment

_____ oval plate
Figure. 2. Ventral view of Laccophilus m. maculosus male.

On occasion the smaller proximus has been confused with maculosus, but the
pattern contrast is much stronger in the latter. The three are similar, however,
in that they all have the same general elytral pattern outline, belong to those
species with a metacoxal file, and have a short prosternal process. No other
Laccophilus from northeastern North America are irrorated.

NOMENCLATURAL NOTES. — Laccophilus m. maculosus has
been ascribed to Germar because Say’s description of maculosus was
long considered to have been published in 1825. LeConte (1891,
p. 435), however, gives the date as 1823 with following footnote,
“The title page of the 2nd vol. of the New Series of these Transactions
bears date 1825, which was the time of completion of the volume, but
the late Dr. T. W. Harris informed me in a letter, that he received
from Mr. Say a copy of this paper, with the following addition to the
title; — ‘Printed and published by Abraham Small, 1823.’ This, of
course, gives Say’s name precedence over those published by Germar
in his Sp. Ins. Nov. in 1825.” Blatchley (1910) did ascribe macu-

MEM. AMER. ENT. SOC., 26

36 THE GENUS LACCOPHILUS

losus to Say. Zimmermann (1920) made reference to LeConte’s edited
work, but either overlooked or ignored his opinion and made no ref-
erence to Blatchley. I have not seen Germar’s type and Say’s are
presumed lost; hence, a neotype is designated.

Say’s description separated two varieties; one (Var. A) is proba-
bly m. maculosus as it is widely recognized today, but the second
(Var. B) is undoubtedly L. fasciatus rufus Melsheimer. It seems
best to accept the first variety as maculosus to avoid more nomencla-
tural confusion.

VARIATION. — This race shows remarkable uniformity. There
is about a 16 percent range in variation in each of four characters
with no clearly evident clinal or other geographic differentiation. For
example, nearby localities in Michigan may have mean values which
differ as greatly as a sample from several hundred miles away in
Iowa (Table 1). Mean size of males is slightly larger than females
(1.8% ), but there is sufficient overlap that small samples usually have
broadly overlapping 95 percent confidence limits. Size overlap is
nearly complete in the two sexes; in 642 specimens (334 females, 318
males) only eight males were larger in total length than any females,
and only three females were smaller than any males (fig. 4).

The elytral pattern reflects the statistical data. The pattern is ir-
regular in outline and intensity of color, but it has much the same
appearance throughout its range. Individual variation appears greater
than regional variation.

There are differences in the male coxal file between maculosus and
decipiens in coarseness and number of lines, but only estimates of the
number of lines are possible. More exact comparisons will have to
wait for auditory analysis. L.m. maculosus has about 25 to 30 lines
in the file and decipiens has about 45.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L.m. maculosus occurs from Alabama to Nova Scotia westward
to northwestern Colorado, western Nebraska, the Dakotas, and Mani-
toba, where it intergrades with L. m. decipiens. It probably occurs
continuously through southern Canada to near Winnipeg. It extends
along the Appalachians to near Atlanta, Georgia. There are single
specimens from coastal South Carolina, southeastern Alabama, and
Dallas, Texas, but the range is probably restricted to near a line run-
ning from-northeastern Georgia to southwestern Nebraska. I have

JAMES R. ZIMMERMAN 37

LACCOPHILUS M. MACULOSUS
LACCOPHILUS M. DECIPIENS °
LACCOPHILUS M. SHERMANI 2
L.M.MACULOSUS X L.M. DECIPIENS 9
L.M. DECIPIENS X L. M. SHERMANI

Figure 3. Distribution of Laccophilus maculosus.

made over two hundred collections in Texas, Oklahoma, and Kansas,
and have collected it only as far south as Reno County, Kansas, which
is about the limit of the range. Single individuals in this and other
species have been collected far out of what might be considered the
normal breeding range. This should be expected for insects that fly
readily and whose adults can survive for short periods in almost any
aquatic situation.

This race prefers shallow, partially shaded ponds that tend to have
some water in them all year. In Indiana they were more common in
glacial, gravelly outwash soils than in clays (Zimmerman, 1960).
They have also been reported in waste-oxidation lagoons in Missouri
(Roberts, Smith and Enns, 1967) in fish hatcheries in Iowa ( Wilson,
O23))r

MEM. AMER. ENT. SOC., 26

38 THE GENUS LACCOPHILUS

MATERIAL EXAMINED

CANADA. — NOVA SCOTIA. Cape Breton Is., 1 ¢, 5 2, viii.19.—, G. K.
MacMillan (CNG); Truro, 1, viii.5.13, R. Matheson (CNL). ONTARIO.
Guelph, 8 (USNM); 2 6, 1 @, iv.20.32, R. H. Ozburn, 2 ¢, vii—.01 (GLPH).
Grimsby, 1 6, 1 2, J. Pettit (GLPH). London, 1 2, 2 4, W. Saunders
(GLPH). Point Pelee N. P., 1 2, v.28.44, B. Malkin (FM). Brimley, 1 2,
vi.28.36 (SCH). Ridgeway, 4, viii.7.89, E. P. Van Duzee (BERK); 1 6, A. H.
Kilman; 1 6, 3 2 (GLPH). Toronto, 2, R. J. Crew (CNL); 1, ix.14.95
(USNM); 1 6, xi.33.—, H. B. Leech (MCHS). QUEBEC. Duparquet, 1,
ix.10.34; 1, viii.20.36; 1, v.15.43; 1, v.15.43; 1, vii.22.43; 1, 1x.2.43; 1, 1x.28.43,
G. S. Smith (CAS); 1 2, vii.1.44, G. S. Smith (FM). Levis, 1, viii.31.35, D.
Dunavan (USNM). Montreal, 2 6, 2 2, vi5.17; 1 6,3 2, vii.1.23, J. Quellet
(MCHS); 2 6, 3 2, ix.—.99 (FM). Quebec, 1 2, v.22.56 (SCH).

UNITED STATES OF AMERICA. — ALABAMA. Russel County. Seale,
1 6, vi.18.54, R. L. Fischer & D. L. Haynes (MCHS). CONNECTICUT.
Fairfield County. Sheffield Island, 2 4,1 2, viii.14.01, J. L. Zabriskic (AMNH).
Stamford, 1 ¢, v—.—, Angel (AMNH). County, 5 ¢, 7 2, viii.S.—, C. H.
Roberts; 1 9, vii.4— (AMNH). Litchfield County. Cornwall, 8, ix.23.20,
1, ix.8.20, 2, x.10.36, Chamberlain (CNL). Goshen, 1 @, vii.5.25, L. B.
Woodruff (AMNH). Litchfield, 1 2, v.1.22; 1 6, 3 2, ix.3.24, L. B. Wood-
ruff (AMNH). Sharon, 1 ¢, 1 2 (LACM). DELAWARE. Kent County.
Bombay Hook, 2, vi.12.54; 11, vi.8-21.54; P. J. Spangler (USNM). _ DIS-
TRICT OF COLUMBIA. Piney Branch, 1, iv.2.05; 2, v.2-16.05; 1, vii.26.06;
1, iv.25.05, D. H. Clemons (USNM). Rock Creek, 1, viii.31.06, D. H.
Clemons (USNM). U. S. Fish Comm., Carp Ponds, 4, vil.21.97 (USNM).
Washington, 1, iii.4.07, W. L. McAtee; 1, xi—.—, 1, ix.13.06 (USNM); 2 9,
iv.15.—, x.20.—, H. S. Barber (USNM); 1 ¢, iv.15.—, H. A. Barber (FM);
1 9, C. H. Roberts (AMNH). GEORGIA. dHall County. Gainesville, 10,
iv.2.11 (CNL); 2, iv.2.11 (USNM). Rabun County. 1 6, 1 9, vii——,
C. W. Leng (CAS). ILLINOIS. Alexander County. Olive Branch, 1 °,
x.8.09, Gerhard (FM). Champaign County. Champaign, 1 ¢, 2 9, xi.5.27,
A. T. McClay (DAV). Cook County. Chicago, 3, ix.17.11, E. Lijablad
(USNM); 2, iii.9.—, H. Soltau (USNM); 2 6, 1 9, iv.26.25; 1 2, vii.22.06,
W. J. Gerhard (FM). Cook Co., 1 6, 1 2, E. B. Chopa (FM). Edge Brook
(Morton Grove), 4, viii.16.04, J. D. Sherman, Jr. (USNM). Evanston, 3 ¢6,
3 2, x.23.04, A. B. Wolcott (FM). South Chicago, 4 6, 4 2, ix.14.02; 1 9,
v.20.06, W. J. Gerhard (FM). McHenry County. Cary, 1 2, vili.27.05, W. J.
Gerhard (FM). Pike County. Pittsfield, 1 3, vi.17.52, B. Cadwell (DAV).
Vermillion County. Muncie, 2, iv.20.07 (USNM). Oakwood, 2 2, v.5.28,
A. T. McClay (DAV). Rossville, 1, vii.10.56, P. S. Spangler (USNM). INDI-
ANA. Brown County. Brown Co. St. Pk., 9 6, 4 2, iv.21.56, JRZ (NMSU).
Clinton County. Kirklin, 3 m. W., 1 ¢, 2 9, ix.12.58; 1 2, ix.9.58;9 6, 6 Q,
x.3.58;J8 6,2 9, x.10.58;9 6, 3 2, x.17.58, IRZ (NMSU). Dubois County.
County, 3 6,5 9, x.29.55, J. Hanegan & JRZ (NMSU). Grant County. Deer
Creek, 5 6, 5 9, iv.2.56, JRZ (NMSU). Hamilton County. Westfield, 1 ¢,

JAMES R. ZIMMERMAN 39

1 2, v.4.56, JRZ (NMSU). Huntington County. County, 1, iv.2.56, JRZ
(NMSU). La Grange County. Eddy, 3 m. W., 1, v.2.56, JRZ (NMSU).
Lake County. Pine (near Gary), 2, (USNM). 2 4, 3 2, v.21.06, A. B. Wol-
cott (FM). Laporte County. Wanatah, 12 ¢, 15 9, viii.12.52, P. J. Spangler
(USNM). Laporte, 1 é (CNG). Kosciosko County. Winona Lake, 2 2,
vii.8.55, JRZ (NMSU). Madison County. Anderson, 7 6,7 92, vi.21.57, JRZ
(NMSU). Marshall County. Myers Lake, 1, v.4.56, JRZ (NMSU). Monroe
County. (Note: Only one record is given here, but more than 500 specimens
taken over a three year period from all parts of Monroe County are in the
author’s collection.) Bloomington, 5m. N., 11 4, 17 2, ix.17.56, JRZ (NMSU).
Morgan County. Waverly, 3 6,5 92, vi.17.57, JRZ (NMSU). Noble County.
Wolflake, 5 m. S., 4 6, 4 2, x.26.56, JRZ (NMSU). Owen County. Rich-
land Creek, 3 6, 7 2, vii.26.55; 3 6, 2 2, x.22.55, JRZ (NMSU). Tippe-
canoe County. County, 1 6, 4 9, 1x.27.58, N. M. Downie (DAV). Vigo
County. Terre Haute, 1, Blatchley (USNM). Wabash County. Wabash,
3m.N., 3 6,3 2, x.26.56, JRZ (NMSU). Wayne County. Richmond, 1 6,
1 92, viii.7.54, PSB (CAS). Whitley County. County, 3 6, iv.2.56, JRZ
(NMSU) . IOWA. Dickenson County. L. Okoboji, 1, vi.27.16; 1, vii.14.16,
Buchanan (USNM). Hamilton County. Jewell, 1 ¢, v.12.41, Sigley (DAV).
Little Wall L., 3 6, 3 9, viii.19.52, P. J. Spangler (USNM). Henry County.
Mt. Pleasant, 1, iv.14.26, Latta (USNM). Mt. Pleasant, 1, iv.29.26 (USNM).
Johnson County. lowa City, 1, iv.7— (USNM). Solon, 1, iv.20.15, L.
Buchanan (USNM). Lee County. Ft. Madison, 1 2 (FM). Story County.
Ames, 1, (CNL); 1, xi.20.32, F. Andre (USNM); 1 9, iv.23.18, H. A. Scullen
(ORES). Indianola, L. Ahquahbi, 4 6, 7 2, vi.14.52, P. J. Spangler (USNM).
KANSAS. Butler County. Santa Fe Lake, 3 6, 2 2, iv.4.58, IRZ (NMSU).
Cherokee County. Galena, 3 (USNM); 1 2, Ebb. Crum (AMNH). _ Dicken-
son County. Elmo, 3 m. S., 1 9, vi.16.58, JRZ (NMSU). Douglas County.
Scorpion Hill, 4 ¢, iv.25.52, P. J. Spangler (USNM). Douglas County, 1 2,
x.9.45, J. Rozen and B. Lindner (BERK). Johnson County. Sunflower, 1 ¢,
1 2, ix.7.52, Spangler and Bell (USNM). Leavenworth County. Leaven-
worth St. Pk., Tonganoxie L., 13 6, 8 2, vi.l.52; 4 6, 11 2, ix.14.52, P. J.
Spangler (USNM). MAINE. Cumberland County. Casco, 1, ix.1-3.45, J. C.
Bradley (CNL). Portland, 2, ix.14.—, J. D. Sherman, Jr. (USNM); 1 4,
ix.11— (AMNH). Kennebec County. 1, G. M. Greene (USNM). Mon-
mouth (Mommouth, sic!), 12, G. M. Greene (ANSP). Oxford County.
Paris, 1, vi.16.44, C. A. Frost (CAS). Reno County. Haven, 4 m. S., 1 2,
vii.19.60, JRZ (NMSU). York County. Kittery Point, Cutts. I., 3, vii—.24
(USNM). Moody, Moody Beach, 1 4, viii.23.54, P. J. Spangler (USNM).
Saco, 2, v.19.38, W. Nutting (ARI). MARYLAND. Baltimore County. Lu-
therville, 1, vii.9.74 (USNM). 3, viii.22— (USNM). Harford County.
Edgewood, 2, ix.2-17.18, H. Dietrich (CNL). Montgomery County. Plum-
mer’s I., 2, iii.24.02, W. L. McAtee; 4, ii.23-29.07, D. H. Clemons; 1, iv.24.07,
W. L. McAtee; 2, H. S. Barber (USNM). Kensington, 22, vi.6-17.61, J. W.
Fitzgerald (USNM). Cabin John, 1, vii.13.—, L. L. Buchanan (USNM).

MEM. AMER. ENT. SOC., 26

40 THE GENUS LACCOPHILUS

Prince Georges County. College Park, 2, x.—.33, R. A. Littleford (USNM).
Hyattsville, 1, vii.4.11 (USNM). Branchville, 2, iv.25.26, L. Buchanan
(USNM). MASSACHUSETTS. Barnstable, Woods Hole, 1, ix.25.81 (USNM).
Berkshire County. Egremont, 2 ¢, 5 2, vi.21.—, C. H. Roberts (AMNH).
Bristol County. Taunton, 1 9, ix.24.36 (AMNH). Essex County. Ipswich,
1 2, vii.l.— (AMNH). Lawrence, 4, ix.25.20, E. W. Mank (CNL); 1,
.19, E. W. Mank (CNL). Hampden County. Springfield, 8, vi.15-17.16;
3, iii.24-28.17, E. A. & W. H. Chapin (USNM); 1, iv.19.00, F. Knat (USNM).
Wilbraham, 1 (CNL); 1, viii—.95 (USNM). Hampshire County. Amherst,
12,1 4, vii.11.26, K. A. Salman (LACM); 1 6, x.4.63, D. April (SCH).
Holyoke, Holly Ledge, 4, iv.24.99, G. Dimwock (CNL). Mt. Tom, 4, iii.25.17,
E. A. Chapin (USNM). Middlesex County. Cambridge, 1 (USNM). Mel-
rose (High), 12, iii.15.—, D. H. Clemons (USNM). Watertown, 1 9, v.17.21,
R. F. Hussey (MCHS). Norfolk County. Milton, 3 6, 2 9, i1i.13.42, H.
Clench & K. Christiansen (CNG); 1 9, iv.3.41, H. Clench (SCH). Needham,
1, ix.1-2.45, J. C. Bradley (CNL). Suffolk County. Brookline, 2 (BERK).
Worcester County. Southboro, 1 6, iv.13.39, J. G. Thorndike (AMNH).
MICHIGAN. Alcona County. Barton City, 4 6, 1 2, vii.23.63, R. B. Willson
(MCHS). Alger County. Chatham, 1 2°, V. E. Shelford (AMNH). Munis-
ing, 10 m. S., Big Twin Lake, 2 ¢, 2 9, vii.22.64, R. B. Willson (MCHS).
Allegan County. Allegan St. Forest, 1 Q, vii.22.63, G.C. Eickwort (MCHS).
Baraga County. Anse, 8 m. S., 2 6,3 @, vii.10-11.64, R. B. Willson (MCHS).
Barry County. County, 1 6, iv.29.55, R. L. Fischer (MCHS). Bay County.
County, 5 6, 1 9, viii.11.51, R. R. Dreisbach (MCHS). Berrien County.
Napier, Wolk Creek, 1 4, ix.17.64, R. B. Willson (MCHS). Near Sawyer,
Warren Dunes St. Pk., 19 6, 23 9, P. J. Spangler (USNM). Stevensville,
2 4, 1 Q, vii.12.63, R. B. Willson (MCHS). Calhoun County. Olivet, 4 m.
S., Indian Ck., 20 6, 20 2, x.5.63, R. B. Willson, R. W. Matthews (MCHS).
Cheyboygan County. Black L., 1 ¢, vi.30.47, P. A. Hawkins (MCHS); 1 6,
vi.23.49, J. D. Lattin (ORES). Douglas L., 3 6, 3 2, vii.4.31, C. W. S.
(MCHS); 5 6, 4 2, vi.24.52, P. J. Spangler (USNM). Nelson L., 3 2,
vili.16.52, J. P. Spangler (USNM). Smith’s Bog, 1 °, viii.4.52, P. J. Spangler
(USNM). Chippewa County. McCarron, Charlotte R., Beaver Ck., 40 6,
41 92, viii.2-4.64, R. B. Willson (MCHS). Clare County. County, 3 ¢, x.5.49,
vili.2.51, R. R. Dreisbach (MCHS). Clinton County. Bath, 1 6, 4 2, iv.18.64,
R. B. Willson (MCHS); 1 3, iv.29.60, C. A. Scheibner (MCHS). Park Lake,
18 6, 21 Q, viil.17.63, R. B. Willson (MCHS). Delta County. Escanaba, 1,
vil.17.—, Hubbard & Schwarz (USNM). Escanaba, Portage Ck., 4 6, 2 2,
R. B. Willson (MCHS). St. James, 17 m. N., 1 2, vi.22.64, R. B. Willson
(MCHS). Eaton County. Grand Ledge, 1, Hubbard & Schwarz (USNM).
Eaton Rapids, 1 4, viii.3.43, C. W. Sabrosky (MCHS). Olivet, Indian Ck.,
2 6,2 2, x.5.63, R. B. Willson (MCHS). Emmett County. Levering, Carp
R:, 1 9) vii.5.52;, Ps J. Spangler, (USNM).) Pellston)) 12067) 72s y evil 52
vili.5-8.52, P. J. Spangler (USNM). County, 1, vii.25.52, P. J. Spangler
(USNM). Genesee County. Flint, 16 8, 12 2, ix.7.63, v.30.64, vii.25.64,
R. B. Willson (MCHS). Gladwin County. White Star, 1 ¢, 1x.4.49, R. L.

JAMES R. ZIMMERMAN 41

Fischer (MCHS). County, 2 6, 1 9, vii.20.49, R. R. Dreisbach (MICHSU).
Gogebic County. Wakefield, Black R., 1 2, vi.17.64, R. B. Willson (MCHS).
Gratiot County. County, 1 6, 3 9, vi.10.49, R. R. Dreisbach (MCHS).
Houghton County. Chassell, 1 6, 1 92, vi.24.64, R. B. Willson (MCHS).
Houghton, 1 ¢, vii.7.64, R. B. Willson (MCHS). Ingham County. E. Lansing,
1 6,1 Q, iv.23.63, 1 @, viii.26.63, 1 2, v.5.38, R. B. Willson (MCHS). Has-
lett, 2 6, ix.13.55, R. Fischer (MCHS). Isabella County, 3 4, 3 2, ix.11.48,
R. R. Dreisbach (MCHS). Kalamazoo County. Gull Lake Biol. Sta., 1 4,
1 2, vii.18.63, Fischer & Matthews (MCHS); 1 ¢, x.28.63, R. B. Willson
(MCHS). Kent County. Grand Rapids, 2 ¢, 2 2, viii.24.37 (MCHS); 2 ¢,
2 2 (FM). Keweenaw County. Anmeek, 3 m. N., 1 é, vii.8.64, R. B. Will-
son (MCHS). Bete Grise Bay, 7 3, 3 2, vii.7.64, R. B. Willson (MCHS).
Isle Royale, 2 6, 1 2, vii.3.64, R. B. Willson (MCHS). Leelanau County.
Glen Arbor, Sleeping Bear Dunes, Glenh., 3 ¢, 3 92, viii.17.52, P. J. Spangler
(USNM). Luce County. Dollarville, 1 ¢, 4 2, vii.31.64, R. B. Willson
(MCHS). Paradiseville, 11 m. WNW., 1 2, vii.31.64, R. B. Willson (MCHS).
Newberry, Tanguamonon R., 4 6, 3 2 (MCHS). Mackinac County. Cedar-
ville, 2 6, 4 2, viii.5.64, R. B. Willson (MCHS). Hessel, Law Ck., 1 6,
vili.5.64, R. B. Willson (MCHS). Marquette County. Champion, 1 9, vii.13.65,
R. B. Willson (MCHS). County, 2; 4, vii.21.6, J. D. Sherman, Jr. (USNM);
8 6, 2 2, vii.17-19.65, R. B. Willson (MCHS). Mason County. Pere Mar-
quette & Great Sable R., 6 6, 8 9, viii.8.52, P. J. Spangler (USNM). Midland
County, 9 6, 7 &, viii.8.48, ix.10.48, x.1.49, vii.25.51, viii.S.51, R. R. Dreis-
bach (MCHS). Missaukee County. Merritt, 12 m. NE., 3 6, 6 2, ix.6.63,
R. B. Willson (MCHS). Monroe County. Monroe, 2, ix.—.—, Hubbard &
Schwarz (USNM). Montcalm County. Flat River Game Area, 2 2, v.14.55,
R. L. Fischer (MCHS). Oakland County. Ortonville, 2 6, 1 2, viii.4.10,
vii.3.28, A. W. Andrews (MCHS); 1 @, vii.14.49, R. R. Dreisbach (MCHS).
Ottawa County. Allendale, 5 ¢, 6 2, vii.17.63, R. B. Willson (MCHS). Zee-
land, 11 6, 14 9, vili.9.63, R. B. Willson (MCHS). County, 2, viiil.31.—,
E. A. P. (USNM). Roscommon County, 1 2, vi.20.53, R. R. Dreisbach
(MCHS). Schoolcraft County. Germfask, 15 6, 32 @, vii.25-29.64, R. B.
Willson (MCHS). Shiawassee County. Owosso, 1 4, vii.23.53, Bruce Wilson
(MCHS). St. Clair County. Port Huron, 1 6, 1 2, vii—.42, D. G. Kelley
(CAS). Van Buren County. South Haven, 1, viii.11.04, J. D. Sherman, Jr.
(USNM); 2 6, 1 2, vi.1.91 (MCHS). Washtenau County. Chelsea, 7 ¢,
1 2, vii.15.54, R. L. Fischer (MCHS). County, 1 6, 1 2, v.11.44, Wilmordk
(CAS); 1 2, iv.18.45, R. L. Fischer (MCHS). Wayne County. Detroit, 2,
vi.l.—, Hubbard & Schwarz (USNM). MINNESOTA. Anoka County, 1,
iv.25.36, H. R. Dodge (CAS). Hennepin County. Excelsior, 1, J. D. Sher-
man, Jr. (USNM). Minneapolis, 1, ix.7.15; 1, vi.29.16; 1, v.25.20, F. C.
Fletcher (CNL). Minnetonka, 4, viii.2.23, F. C. Fletcher (CNL). Itasca
County. Little Winnebegosish (Lake?), vi.10-11.35, K. Cooper (USNM).
Olmsted County. Rochester, 10, J. D. Sherman, Jr. (USNM). Ottertail
County. Fergus Falls, 1, viii.19.11, Stoner (USNM). Ramsey County. St.

MEM. AMER. ENT. SOC., 26

42 THE GENUS LACCOPHILUS

Paul, 5, J. D. Sherman, Jr. (USNM). Renville County. Buffalo L., 10, J. D.
Sherman, Jr. (USNM). Rock County. Lucerne, 1 2, vi.14.55, W. A. Drew
(MICHSU). MISSOURI. Barry County. Cassville, Roaring River St. Pk.,
1, iv.22.55, M. C. Grabeau (USNM). 1 6, x.8.54, P. J. Spangler (USNM).
Boone County. Ashland, 2 6, 3 2, v.8.55, P. J. Spangler (USNM). Columbia,
1, v.8.37, W. M. Gordon (CNL). 1, vi——, 2, C. V. Riley (USNM). 1 9,
x.8.54, P. J. Spangler (USNM). Deer Pk., 6 m. E., 1, viii.28.53, P. J. Spangler
(USNM). Callaway County. Kingdom City, 4 m. N., 1, vii.16.53, P. J.
Spangler (USNM). Crawford County. Meramec R., St. Rd. 8, 3, vii.21.53,
P. J. Spangler (USNM). Lawrence County. Verona, 1 46, vii.7.53, P. J.
Spangler (USNM). Shannon County. Winona, Lewis L., 3 6, 4 2, iv.9.55,
P. J. Spangler (USNM). St. Louis County. Howard Bend, 1, vi.28.37, W. M.
Gordon (CNL). St. Louis, 4, vii.—.77 (USNM); 1 92, vi.8.32 (CRNG).
Vernon County. Near Nevada, 2 ¢, 1 9, v.14.58, J. W. McReynolds (CAS).
NEBRASKA. dAolt County. Stuart, 2m. NW., 1, vi.3.37, C. E. Burt (USNM).
Lancaster County. Lincoln, 7 6, 6 2, x.15.60, W. F. Rapp, Jr. (NMSU).
Logan County. Stapleton, 1 m. N., 2 4, vii.15.67, H. B. Leech (CAS). Holt
County. Stuart, 2 m. NW., 1, vi.3.37, C. E. Burt (USNM). Otoe County.
Palmyra, 1 6, W. F. Rapp, Jr. (WFR). NEW HAMPSHIRE. Carroll County.
N. Conway, 1, vi.15.46, C. A. Frost (CAS). Coos County. Carter Lake, 12
(USNM). Jefferson Junction, 1, J. D. S., Jr. (USNM). Mt. Adams, 4 6,
vul.24.—; 1 92, vii.l4.—; 7 6, 2 9, vill.27— (AMNH). Mt. Washington, 7;
2, vii.30.95, J. D. S., Jr. (USNM):; 2 6, 32, Mrs. A. T. Slosson; 2 2, ix.m—.—,
J. D. Sherman, Jr.; 1 2, vi.—28, R. Dow (USNM). Pittsburg, Conn. R., 6,
vill.16.25, Chamberlain (CNL). Randolph, 6, J. D. Sherman, Jr. (USNM).
Starr Lake, near Mt. Adams, 55; 1 2, J. D. Sherman, Jr. (AMNH). White
Mts., 1 2 (AMNH). NEW JERSEY. Bergen County. Dumont Woods, 1 2,
v.6.34, C. L. Ragot (FM). Passaic R., 4, vii.1.33, Shoemaker (USNM).
Ramsey, 4 6, 2 2, vi.7.—, C. H. Roberts; 2 ¢, 3 2, vii.13-16.12 (AMNH).
Westwood, 107, ix.7.—; 2 6, 4 2, vii.26.—;4 6,5 Q, viii.30.—; 1 4, vii.7.—;
5¢6,72,x.11—;44, 192, x.17— (AMNH). County, 1, iv.20.02; 4, v.11.02,
Shoemaker (USNM). Burlington County. Bridgeboro, 1 6, iv.26.41, E. J. F.
Marx (AMNH). Hartford, 1 4, 1 2, xii.28.40, E. J. F. Marx (AMNH).
Medford, 2 6, vii.20.41, E. J. F. Marx (AMNH). Moorestown, 1 6, 1 2,
vi.29.41, E. J. F. Marx (AMNH). Lanola, 9 6, 12 2, vi.29.41; 2 4, viii.17.40,
E. J. F. Marx (AMNH). Camden County. Clemonton, 12, vii.26.96; 1,
v.16.97; G. M. Greene (USNM); 1 4, vii.l.— (ANSP). Delair, 1 ¢, vi.l.—
(AMNH). County, 1 6, vi.8.—, E. L. Dickerson (AMNH). Essex County.
Newark, 3 ¢, E. L. Dickerson (AMNH). Gloucester County. Pitman, 1,
ix.22.37, J. C. Bradley (CNL). Middlesex County. Avenel, 2 6, v.16.—,
Berk (MCHS); 1 2, v.22.31, Siepmann (FM). Amboy Mdw., 2 6, vi.27.—,
E. L. Dickerson (AMNH). Spotswood, 1 6, vii.4.—, E. L. Dickerson (AMNH).
Morris County. Boonton, 2, v.10-12.01; 1, iii.30.17, G. M. Greene (USNM).
Lincoln Park, 1, viii.13.02, R. Godfrey (USNM). Lower Longwood.?, Rocka-
way R., 1 2 (BERK). Ocean County. Lakehurst, 1, ix.1.01; 1 (USNM);
3 6,2 2, v.24.—3; 1 2, v.31.—; 2 ?, vi.13.—, C. H. Roberts (AMNH). Pas-

JAMES R. ZIMMERMAN 43

saic County. Patterson, 1 6, iv.21.—; 1 4, v.3.—; 4 6, 1 9, vii.4— (AMNH).
Wanague, 2 6,1 9, v.12.40, B. Malkin (FM). Somerset County. Metuchem,
1 6, v.5.— (AMNH). Milltown, 1 6, 1 2, ili.14— (AMNH). New Bruns-
wick, 1 2, vii.27.—; 1 6, vill.9.—, E. L. Dickerson (AMNH). Union County.
Rahway, 1 ¢, vii.19.31, Siepmann (FM). 1 6, iv.20.—, E. L. Dickerson
(AMNH). Warren County. Great Piece? Mew., 3 4, ix.3.—, E. L. Dicker-
son (AMNH). Hope, 1 ¢, 1 9, xi.9.41, E. J. F. Marx (CAS). NEW YORK.
Bronx County. Bronx Pk., 1 ¢, 3 2, viii—m —; 21, ix—.— (AMNH). Van
Cordlandt Pk., 1, ix.13.—, Shoemaker (USNM); 1 4, v.4.41, Greame Kelley
(CAS); 1 4, v.4.41 (FM). Chataqua County. L. Chataqua, 1 ¢, z 2,
viil—.— (CNG). Courtland County. McLean Bogs, 10, viii.7.17, H. Diet-
rich (CNL). Mud Pond, McLean Res., 1, ix.24.25 (CNL). Erie County.
Buffalo, 2 (USNM); 3 2 (AMNM); 3, E. P. Van Duzee (BERK). Colden,
1 2, vii.33.16, M. C. Van Duzee (CAS). Evans, 1, vi.16.85, C. V. Riley
(USNM). Lancaster, 10 6, 1 9, vil.20.08, M. C. Van Duzee (CAS). Tona-
wanda, 1 Q, vii.9.16, M. C. Wan Duzee (CAS). Essex County. Au Sable
Lake, 3 (USNM). Mt. Marsey, L. Tear, 1, J. D. Sherman, Jr. (USNM). Mt.
Mary, 1, viii.6.14, J. D. Sherman, Jr. (USNM). Franklin County. Coreys,
1 9, x.12.28, W. J. Gerhard (FM). Green County, 12 (CNG). Kings
County. Cypress Hills, 3 2, v.17.90; 1 6, 1 6, v.21.90; 2 6, 1 2, v.23.90;
1 @, vi.5.90; 4 9, viii.17.91, J. L. Z. (AMNH). Flatbush Waterworks, 4 ¢,
5 6, iv.30.90; 1 2, v.3.90; 1 9, v.12.90 (AMNH). Manhattan County.
N.Y.C., 5, Hanwitta (USNM). Monroe County. Rochester, 2 9, vii.1.42;
18 6, 10 9, ix.9.44; 1 6,1 @, ix.23.44; 2 6, 1 9, vi.23.45; 1 6, 1 9, vi.30.45,
F. C. Fletcher (LACM). Nassau County. Wantaugh, L. I., 3 6, vii.22.39;
3 6, 1 9, iv.12.41 (FM). Niagara County. Olcott, 1, iv.3.21; 1, vii.17.21;
4, vili.14.27, H. Dietrich (CNL). Onondaga County. Manlius, 2, ix.4.71,
H. H. Smith (CNL). Orange County. Ft. Montgomery, 1 6, iv.20.41 (FM).
West Point, 4, iv.19.09; 1, v.21.10; 14, vi.9-18.10, W. Robinson (USNM).
Cunningham Park, 2 4, vii.13.41; 2 2, ix.30.40-41, G. Kelley (CAS). Forest
Pk., 1, v.15.04; 4, x.23.04; 2, v.16.i0, Shoemaker (USNM). Kissena Park,
2 46, 1 2, vili.30.40, Greame Kelley (CAS). Richmond County. Staten Is-
land, 4, x.23.04; 1, iv.12.08; 3, iv.16.05 (USNM); 2 6, 6 92, ix.27.38: 3 6,
PCE RX eS SAG epiVal ESO slo 4 Oemiv- Ors Os 1) Om val4i39 0) 6), 4 OR v.20:39:
1 6, vi.7.39; 1 2, ix.9.39; 1 9, vii.22.40 (FM); 1 3, 2 9, iv.19.—, C. H. Rob-
erts; 1 6, v.7.25, Woodruff; 3 6 (AMNH). Tompkins Cave, 9 6, 16 2,
vili.10.10 (AMNH). Rockland County. Nyack, 5 é6, 4 2, x.—.86, J. O. Za-
briskic (AMNH). Scarsdale, 1 ¢, v.4.20 (MCHS). St. Lawrence County.
Canton, 1 6,1 2, v.15-16.33 (ORES). Schuyler County. Watkins Glen, 1 ¢,
Mrs. A. T. Slosson (AMNH). Suffolk County. Fisher’s Island, 3 6, 2 2,
vili.17.91, J. L. Z.; 24 3, 13 2, viii.29.90 (AMNH). Orient, L. I., 1, ix.20.42,
Roy Latham (CNL). Tomkins County. Ithaca, 1, vii.26.07; 2, iv.15.13; 1,
ivV.12.15; 4, iv.15.15; 1, iv.25.15; 2, v.15.15; 1, vii.6.16; 9, viii.17.16; 1, viii.23.16;
3, 1x.2.16; 4, ix.10.16; 5, ix.17.16; 2, ix.22.16; 1, ix.24.16; 1, vii.9.17; H. Diet-
rich (CNL); 1, vii.12.20, E. Mank; 1, vii.20.22; 1, viii.17.22; 2, ix.5.22 (CNL);

MEM. AMER. ENT. SOC., 26

44 THE GENUS LACCOPHILUS

4, vii—.— (CNL); 4, Chttn. (USNM). Cascadilla, 1, v.28.24, F. C. Fletcher;
1, iv.28.39; W. M. Gordon; 1, viii.24.40, E. Dietrich (CNL). Ulster County.
Esopus, 2 9, ix.2.—, C. H. Roberts (AMNH). W. Shokan, 11 ¢, 6 2, vii.30.09
(AMNH). Washington County. W. Hebron, 1 ¢, 1 2, vi.30.—, C. W. Leng
(AMNH). Wayne County. Sodus Bay, 1, viii.14.09, A. C. Weed (USNM).
Westchester County. WN. Rochelle, 1 6, viii.3.10 (AMNH). Peekskille, 1,
viii.16.90 (CNL); 7, iv.14.90, J. D. Sherman, Jr. (USNM): 1, x.18.—, J. D.
Sherman, Jr. (USNM). Yonkers, 1, ix.7.—, Shoemaker (USNM). NORTH
CAROLINA. Buncombe County. Asheville, 3, iv.21-26.44, W. E. Hoffman
(USNM); 1 2, ix—95 (AMNH). Black Mts., 4 6, 2 9, v.17.—; 1 4,
vili.19.06, W. Beutenmuller (AMNH). Macon County. Highlands, 1 4, vi.8.61,
O. S. Flint (USNM). Madison County. Hot Springs, 2 2, Mrs. A. T. Slosson
(AMNH). Richmond County. Hoffman, 2 6,3 2 (NMSU). Swain County.
Fontana, L., 2 6, v.18.27; Bryson City, 8 m. SW., 1 2, v.18.57, JRZ (NMSU).
NORTH DAKOTA. Nelson County. Stump, L., E. Gate Pond, 2, vii.7.21,
C. K. Sibley (CNL). Petersburg, 1 2, vii.20.54, PSB (CAS). Richland County.
Hankinson, 1, vii.23,24.17, D. C. Mabbot (USNM). OHIO. Athens County.
Athens, O. U. Airport, 8, vi.20,30.50, P. J. Spangler (USNM); 1, ix.17.50,
P. J. Spangler (USNM). Waterloo Twnp., 2, iv.6.50, P. J. Spangler (USNM);
10, vi.27.50, P. J. Spangler (USNM). Locust Heights Nt. Pt., 2, x.15.49,
P. J. Spangler (USNM). Waterloo Lake, 4, iv.23.50, P. J. Spangler (USNM).
Burr Oak Dam, 2, vii.22.50, P. J. Spangler (USNM); 1 6, ix.30.56, W. C.
Stehr (CAS). Coshocton County. Conesville, 1, vii.23.50, P. J. Spangler
(USNM). Cuyahoga County. Cleveland, 1, vii.—.17, H. Soltau (USNM).
Franklin County. Columbus, 2 6, 2 2, vii.20.40 (AMNH). Fulton County.
Brush Ck., 2 ¢, ix.17.52, I. Slesnick (USNM). Wausson, 5 6, 7 9, ix.21.52,
I. Slesnick (USNM); Harrison L., 5 6, 12 9, ix.27.52, I Slesnick (USNM).
Hamilton County. Cincinnati, 1, vii.17.—, Wickham (USNM). Lytle Creek,
viili.2.51, A. R. Gaufin (AMNH). Lucas County. Cedar Pt., 2, vi.25-26.13,
Jaques (USNM). Muskingum County. Adams Mill, 1, viii.25.50, P. J. Spang-
ler (USNM). Ottawa County. Oak Harbor, 4 6, 1 9, viii.8-9.61, C. P. Mc-
Roy (MCHS). Put-in-Bay, 1 6, 1 2, vii.15.61, C. P. McRoy (MCHS). Ross
County. Londonderry, 2, viii.5.50, P. J. Spangler (USNM). Sandusky County.
Castalia, 1 2, vili.18.61, C. P. McRoy (MCHS). Scioto County. Roosevelt
Lake, 2, vii.29.50, P. J. Spangler (USNM). Vinton County. Lake Hope, 1 ¢,
vil.22.52; 2 6, 2 2, x.7.58, W. C. Stehr (CAS). Warren County. Mason,
2m.S.,6 46, 7 9, vi.24.57, JRZ (NMSU). Wayne County. County, 1, v.21.41,
R. W. Rines (CAS). Killbuck Swamp, 2 é, v.21.41, R. Rines (FM). Wil-
liams County. County, 9 6, 10 9, ix.23.52, I. Slesnick (USNM). Wyandot
County. Sycamore, 5 6, 18 @, viii.10.54; 1 6, 7 2, viii.12.54, P. S. Bartholo-
mew (CAS). PENNSYLVANIA. Adams County. Arendtsville, The Nar-
rows, 3, vii.4.51, P. J. Spangler (USNM). Gettysburg, 1, x.1.—, J. D. Sher-
man, Jr. (USNM)._ Starrers, 5, vi.15.47, P. J. Spangler (USNM). Allegheny
County. County, 3, iv.5.90, E. A. Klages (CNL). County (?), 14,3 2 (FM);
5 6,2 2 (CNG). Harmarville, 2 6, 1 2, ix.l1.— (CNG). Pittsburgh, 2 ¢,
6 ©, ix.4.— (ORES); 11 6,5 2, ix.2.—, 191 stacked on cards, ix.4.—; 19 ¢,

JAMES R. ZIMMERMAN 45

22 2, ix.4.—; 12 6, 11 9; 3 6, 3 Q, vii.14.—; 2 6, 1 9, vii—.02; 2 6, 1 9,
v.8.25; 1 2, v.25; 2 6, 3 2, v.28.—; vii.16.24 (CNG). Wilmerding, 2 ¢,
22 (CNG). Berks County. Kempton, 1 2, vii.12.46, P. Vaurie (AMNH).
Lenhartsville, 2 6, 2 9, vii.10.46, P. Vaurie (AMNH). Bucks County. To-
hickon, 1 9, ix.29.41, E. J. F. Marx (CAS). Clearfield County. Shawville,
1 6, vii.20.40, J. Bauer (CNG). Delaware County. Ardmore, 4 6, 11.14.33;
1 6, 2 2, iv.9.33; 2 6, 2 9, iv.22.33 (SCH). County, 2 6, 1 2, v.18.—
(ANSP). Erie County. Lake Le Boeuf, 2 2, G. E. Wallace (CNG). Presque
Isle, 1 2, vii.29-viii.3.40, G. E. Wallace (CNG). Fayette County. Ohiopyle,
3 6, 6 Q, viii.28.07, Hugo Kahl; 1 2, vi.30.31, W. D. MclIlvoy, Jr. (CNG).
Franklin County. Blue Ridge Summit, 1 é, viii—.43, D. Rockefeller (AMNH).
Chambersburg, 1, J. D. Sherman, Jr. (USNM). Laurel L., 1 6, 3 9, vili.S.—,
P. J. Spangler (USNM). Lackawanna County. Spring Brook, 4, v.22.45
(USNM). Lebanon County. Mt. Gretna, 12, iv.5,9.45, P. J. Spangler (USNM).
Indiantown Gap M. Res., 2, iv.7.50, P. J. Spangler (USNM). Lehigh County.
Lehigh Gap, 1, vii.11.99; 10, vii.12.04; 6, viii.10.04; 79, vii.24-27.05; 6, vili.3.05,
G. M. Greene (USNM). Luzerne County. Dupont, 1, viii.16.04, H. L.
Townes (USNM). Newtown, 1 2, v.16.25, C. B. Woodruff (AMNH). Mon-
roe County. Canadensis, 1, vii.15.26, Shoemaker (USNM). Northampton
County. Easton, 1 ¢, 3 9, vii.21.34, J. W. Green (FM). Perry County.
New Bloomfield, 1 2, vii—.40 (AMNH); 1 2, vii—.40, A. B. Champlain
(LACM). Philadelphia County. Philadelphia, 1, vii.14.—, C. Ilg. (CNL);
4, x.17.—; 1, vii.8.97, G. M. Greene (USNM); Philadelphia Neck, 66, x.30.—;
12, x.17.96, G. M. Greene (USNM); 1, vii.15.04, Kaeber (USNM); 2 ¢, 6 2,
v.29.98 (FM); 1 6 (AMNH); 1 ¢, 2 2, v.4.—; 1 2, v.12.—; 2 6, v.18.—;
1 2, vii.l17.—; 8 6, 3 9, ix.13.— (ANSP). Mt. Airy, 1 6, v.30.— (ANSP).
Pike County. Milford, 3 ¢, v.1.41, B. Malkin (FM). Somerset County.
Kooser St. Pk., 1, viii.11.51, P. J. Spangler (USNM). Washington County.
Canonsburg, 3 6, 1 9, 1.6.52, C. W. Staford; 1 2, iv.6.51 (CNG). West-
morland County. Jeannette, 1 2, vi.—.—; 6 6, 7 2, vii. —; 2 6, vii.4.—;
1 2, vii.10.—; 1 4, vii.l4.—; 1 ¢, vii.15.—; 16,4 9; H. G. Klages (CNG).
York County. Davidsburg Run, 12, vi.11.50, P. J. Spangler (USNM). Sheeps-
head Bridge, 7, iv.7.50, P. J. Spangler (USNM). Washington Twp., 1 6, 2 2,
vii.7.61, P. J. Spangler (USNM). Wrightsville, 1, vii.29.51, P. J. Spangler
(USNM). York, 2 m. SW., 2, iv.10.50, P. J. Spangler (USNM). SOUTH
CAROLINA. Horry County. Little River, 1, xii.15.29, D. Dunavan (USNM).
Pickens County. Pickens, 2, vii.3.27, D. Dunavan (USNM). SOUTH DA-
KOTA. Brookings County. Brookings, 4, J. D. Sherman, Jr. (USNM). Mc-
Pherson County. Leola, 4 m. W., 1 2, vii.24.40, H. C. Severin (CAS). Todd
County. Rosebud, 3 m. E., 1 2, ix.5.40, H. C. Severin (CAS). TENNESSEE.
Campbell County. Cove L. St. Pk.,2 6,1 2, viii.14.50, F. N. Young (AMNH).
Sullivan County. Bristol, 1, v.7.51, B. A. Barrington (CNL); 1 6, 1 2, v.7.51
(AMNH). TEXAS. Dallas County. Dallas, 1 6 (AMNH). VERMONT.
Bennington County. Bennington, 1 (USNM). County, 2 2 (CNG); 4 é,
5 2, vii—.94, C. H. Roberts (AMNH); 1 é (ANSP). Chittenden County.

MEM. AMER. ENT. SOC., 26

46 THE GENUS LACCOPHILUS

Mt. Mansfield, Lake Cloud, 1, vi.29.15 (USNM). Windsor County. Wood-
stock, 1 ¢, vii, F. E. Winters (CAS). VIRGINIA. Fairfax County. Black
Pond, 1, ix.24.15, Shoemaker (USNM). Falls Church, 2, viii.14.—, St. George
& Craighead (USNM). Great Falls, 2, ix.9.06, D. H. Clemons (USNM); 2,
J. D. Sherman, Jr. (USNM); 1, ix.23.15, Shoemaker (USNM). Mt. Vernon,
1, viii.1.19, Shoemaker (USNM). Fauquier County. 1, v.22.61, P. J. Spangler
(USNM). Giles County. Mt. Lake, 1, viii.14.41, A. C. Cole (CAS). Staf-
ford County. Fredericksburg, 3, iii.15.91; 1, ix.—.93; 1, x.1.93; 2, i11.29.99;
1, iv.27.—, Richardson (USNM). WEST VIRGINIA. Greenbriar, White Sul-
phur Springs, 4, vii.23.10, W. Robinson (USNM). Hampshire County. Rom-
ney (near), 1 6, vi—.35 (CNG). Jefferson County. MWHarpers Ferry, 17,
ix.12.17, J. E. Benedict, Jr. (USNM). Mercer County. Athens, 2, vi.16.30,
J. G. Needham (CNL). Preston County. Terra Alta, 4, ix.28.30, J. G. Need-
ham (CNL). Summers County. Talcott, 1 6, vi.5.35 (CNG). Wetzel County.
Lake Floyd, 1, viti.12.50, P. J. Spangler (USNM). WISCONSIN. Bayfield
County. Bayfield, 1 (USNM). Dane County. Madison, 1 92, viii.11.37
(MCHS); 1 4, vi.3.29 (FM); 20+, v.9.36, H. R. Dodge (CAS). Jackson
County. Pigeon Falls, 8 ¢, 4 2, vi.15.52, P. J. Spangler (USNM). Oneida
County. Three Lakes, 1 6, 3 9, W. S. Marshall (AMNH). Sauk County.
Sauk City, 2 (USNM). Shawano County. Shawano County, 20+, vii.17.35,
H. R. Dodge (CAS). Vernon County. Victory, 1, vii.29.27, F. M. Uhler
(USNM). Winnebago County. Oshkosh, 1 é, vii.16.—, E. S. Ross (CAS).

Laccophilus maculosus decipiens LeConte, new status
(Figs. 34, 37, 278, 279)

Laccophilus decipiens LeConte, 1852, p. 205. Type: Museum of Comparative
Zoology, number 5974, male; type locality here restricted to 2.8 miles east
of La Grange, Vizard Creek, Stanislaus County, California; Crotch, 1873,
p. 400; Sharp, 1882, p. 10; Horn, 1894, p. 313; Leech, 1948b, p. 402;
LaRivers, 1951, pp. 397-8; Leech & Chandler, 1956, p. 314; Gordon &
Post, 1965, p. 12.

Laccophilus truncatus Mannerheim, 1853, p. 160.
Laccophilus californicus Motschulsky, 1859, p. 172.
Laecophilus (sic!) maculosus Walker, 1866, p. 317.
Laccophilus fusculus Sharp, 1882a, p. 290.

DIAGNOSIS. — There are only three other irrorated Laccophilus that occur
within the range of L. m. decipiens in western North America. All three,
L. mexicanus, L. fasciatus terminalis, and L. proximus, are smaller than de-
cipiens. Almost all decipiens exceed 5.2 mm which is a maximum length for
any of those three. L. proximus which overlaps decipiens east of the Rocky
Mountains, attains a length of only 4.6 mm. All males and most females of
L. mexicanus have black on the venter instead of brown or reddish-brown as in
decipiens. Some females of mexicanus from coastal California and Oregon
have brown venters, however, and a careful comparison of size and elytral pat-

JAMES R. ZIMMERMAN 47

tern is necessary to separate the two species. L. mexicanus has a more uni-
formly irrorate pattern. JL. f. terminalis, on the other hand, is less evenly
irrorate than decipiens and has a checkered appearance when viewed mega-
scopically. Some depigmented specimens are not checkered, but they are much
paler than decipiens.

L. m. decipiens differs from L. m. maculosus in its larger size (5.1 to 6.4
mm as compared to 4.7 to 5.8 mm), more evenly irrorate pattern, and the
structure of the aedeagus. It is quite similar to L. m. shermani in size, color
and pattern (figs. 276-279), but differs in the more anterior position of the fe-
male epipleural flange, a finer coxal file (about 45 lines to 35), and in the
aedeagus. It is flattened at the tip in decipiens and twisted with a sinus and
incomplete hook at the tip in shermani.

NOMENCLATURAL NOTES. — The synonymy has been con-
firmed except for truncatus Mannerheim and californicus Motschulsky.
Dr. F. N. Young has examined the female type of fusculus Sharp and
has sent me a camera lucida drawing and there is nothing to distin-
guish it from numerous specimens of decipiens. It is probably not
shermani Leech since the epipleura are not expanded; and Nevada,
given as the type locality, seems to be entirely out of the range of that
race. Since the description is based on a single female specimen and
there is no precise locality, it is probably impossible to be certain.
Motschulsky’s californicus was taken at San Francisco, California,
and could conceivably prove to be either L. mexicanus atristernalis
Crotch or L. fasciatus terminalis Sharp. Crotch (1873) had, how-
ever, synonymized truncatus and californicus when he described atri-
sternalis and presumably knew their identity. Mannerheim’s truncatus
was described from Alaska and the Pribilof Islands which is within
the range of biguttatus Kirby and it is remotely possible that it could
be a synonym of that frequently misidentified species. I have exam-
ined LeConte’s type and compared it to a specimen collected by H. B.
Leech from Stanislaus County, California, and could find no differ-
ence other than that of normal individual variation.

VARIATION. — Although exhibiting more variation than L. m.
maculosus, decipiens has relatively little considering its enormous geo-
graphical range and ecological and topographic amplitude. Speci-
mens from near the Coast and in Nevada, however, do appear to be
darker with more suffusion of color between the dots and a more pro-
nounced delineation of the marginal clear areas than those from the
Rocky Mountains and Great Plains. The lateral spots may also be

MEM. AMER. ENT. SOC., 26

48 THE GENUS LACCOPHILUS

smaller. Specimens from Wyoming appear faded when compared to
coastal ones.

The mean length of males is about 5.8 mm and that of females
about 5.6 mm, with an overlap of males and females of about 50 per-
cent. This contrasts with maculosus in which about 75 percent or
more of the males and females overlap. The variation among locali-
ties is slight and an analysis of variance for 240 males from eleven
scattered localities for total length shows no significance at the 95
percent level.

The PW/EL ratio also shows a strong uniformity within decipiens.
Mean values vary from 0.598 to 0.626 for eleven localities. The
ratio does show, however, that decipiens in addition to being slightly
larger than m. maculosus, is also relatively wider. The latter has
mean values ranging from 0.580 to 0.600. The ratio is about the
same for L. m. shermani as decipiens, but the former may be slightly
narrower since there are five samples with values below 0.595 (none
in decipiens).

An attempt was made to count the lines in the metacoxal file of
specimens from different localities. There was no apparent geographi-
cal difference. The highest count was 50, and was found in both
Wyoming and Oregon specimens and the lowest count was 40 in indi-
viduals from Utah and California. The average number was about
44. Only about five to ten percent of the females have epipleural
flanges.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L.m. decipiens (fig. 3) ranges from northern Baja California to
Alaska along the Pacific Coast. It extends eastward to Alberta and
central Colorado. It intergrades with shermani along the northern
edges of New Mexico, Arizona and into eastern Riverside County,
California. There is a single specimen from Hale County in the
northern panhandle of Texas. It occurs from sea level to 9000 feet.
The principal difference between the habitats of decipiens and macu-
losus is that the former is found in or near coniferous forest instead of
temperate deciduous forest. Practically any non-marine aquatic en-
vironment other than large lakes and streams provides suitable habitat
for the adults.

MATERIAL EXAMINED

CANADA. — ALBERTA. Calgary, 1 6, 1 2, vii.4.53; 1 6, 1 2, iv.30.57;
1 6,1 2, v.12.57: 1 &, xii.7.57, F. S. Carr (CARR); 10, x.16.27, O. Bryant

JAMES R. ZIMMERMAN 49

(CAS). Champion, 1 2, v.18.59, F. S. Carr (CARR). Cypress Hills, 1 @,
vi.13.24; 1 9, v.13.28 (DAV). Edmonton, 1 2, v.1.26, F. S. Carr (DAV).
Ft. McCleod, 3 6,5 Q, iii.18.56, F. S. Carr (CARR); 50+, ix.15.28, O. Bryant
(CAS). High River, 1, ix.2.28, O. Bryant (CAS). Medicine Hat, 1 6, 1 92,
x.12.58; 4, viii.25.23, F. S. Carr (CAS); 4, 1x.3.23; 4, viii.7.26, Shoemaker
(USNM). Pincher Ck., 2 2, viii.6.28 (DAV). Porcupine Hills, 1 2, x.18.59,
F. S. Carr (CARR). BRITISH COLUMBIA. Bon Accord, 21, Hubbard &
Schwarz (USNM). Creston, 2, ix.4.54, G. Stace Smith (CNL). Fernie, 10,
ix.16.28, O. Bryant (CAS). Golden, 2 ¢, 1 2, 8.00.03, P. Gregson (AMNH).
Kamloops, 2, ii.1.29, O. Bryant (CAS). Marysville, 1, vi.6.56, E. Fodor
(CNL). Osoyoos, 1, iii.29.41, H. B. Leech (CAS). Victoria Is., Royal Oak,
2, v.19.56, E. Argyle (CNL). Victoria, 1, Wickham (USNM). Vernon, 4,
v.1.29, R. Hopping (ISU). Windermere, 8, ix.17.28, O. Bryant (CAS). Wynn-
del, 10, viii.11.54, G. Stace Smith (CNL).

UNITED STATES OF AMERICA. — ARIZONA. Mohave County. Little-
field, 2 6, 1 2, ix.6.53, P. D. Ashlock (CAS); same date, 1 ¢, 1 2 (DAV).
CALIFORNIA. Alameda County. Albany, 3, iii.16.21 (BERK). Berkeley,
3 6, 9 2, x.15.37, vi.7.37, J. J. Dubois (BERK); 5, x.4-25.19, H. Dietrich
(BERK); 4 ¢, 2 9, xii.14.29, A. T. McClay (DAV); 2, iv.20.41, H. P. Chand-
ler (BERK). Berkeley, Strawberry Pool, 3 2, iii.24.48, R. Johnston (BERK).
Livermore, Arroyo Mucho, 2 6, x.7.56; 16 m. S., 1 2, x.7.56, J. Chemsak
(BERK); 1 2, x.8.54, A. M. Barnes (BERK); 1 2, iii.24.64, W. W. Wade
(CAS); 1 2, viii.7.29, A. T. McClay (DAV). Alpine County. Monitor Pass,
8300 feet, 1 2, vili.24.63, H. B. Leech (CAS). Amador County. Volcano
Cn., 1 6, xi.3.57, R. E. Rice (DAV). Calaveras County. Altaville, 4.5 m. W.,
1 6, vili.31.63, H. B. Leech (CAS). Mokelumne Hill, 1, vii——, Blaisdell
(CNL). Contra Costa County. Tilden Park, 2 2, x.27.48; 1 2, xi.9.48, R.
Johnston, 1 3, xi.21.56, J. A. Chemsak (BERK). Vine Hill, 3, vi.20.10,
Chamberlain (CNL); 1 6, vi.20.10, Chamberlain (AMNH). County, 1 ¢,
vi.3.09, F. W. Nunenmacher (FM). Eldorado County. Camino, 3 m. S., 4 ¢,
6 2, vi.23.48, Carter, Myers, & Smith (BERK); 2 6, 2 2, same date, H. P.
Chandler (DAV). Chile Bar, 3 6, 2 2, vii.5.48, H. P. Chandler (BERK).
China Flat, 2 2, vii.15.48 (BERK). Echo Lake, 3, vii.13.32, E. O. Essig
(BERK). Pollock Pines, 1 ¢, 2 9, vi.29.38; 1 92, vi.24.38; 1 6, vii.3.36; 1 6,
5 2, vi.23.39; 1 $, 1 2, vii.8.39; 1 3, viii.18.39; 1 3, vii.23.39; 1 9, ix.1.39;
1 3, 3 9, vii.8.48, A. T. McClay (DAV). Rubicon R. (near Georgetown),
1 2, vii.28.63, H. B. Leech (CAS). Tahoe, 1, vii.3.50, R. Schuster (BERK).
County, 1 2, vi.8.06, F. W. Nunenmacher (FM). Del Norte County. Smith R.
at Hwy. 101, 1 6, ix.29.65, H. B. Leech (CAS). Fresno County. Fresno, 1,
v.30.—, F. A. Schwarz (USNM). Wartham Ck., near Parkfield, 1 6, xi.30.63,
H. B. Leech (CAS). Humboldt County. Ferndale, 1, ix.14.46, H. P. Chandler
(BERK). Bull Ck. at Cuneo Ck., 1 2, x.2.65, H. B. Leech (CAS). Inyo
County. Deep Springs, 1 ¢, vi.19.54, P. Raven (CAS). Keeler, 1, vii.6-14.—,
Wickham (USNM). Lone Pine, Owens Valley, 1, iv.20.31, Blackwelder
(USNM); 1 6, 2 2, v.20.37, J. J. Dubois (BERK). Olancha (?) 1, J. D.

MEM. AMER. ENT. SOC., 26

50 THE GENUS LACCOPHILUS

Sherman, Jr. (USNM). Kern County. Red Canyon, 1 2, vi.10.22 (AMNH).
Lake County. Thurston Lake, 1, H. P. Chandler (BERK). Lassen County.
Eagle Lake, 1, ix.19.46, H. P. Chandler (BERK). Hallelujah-Jct., 1 9, vii.4.49,
J. W. McSwain (BERK); 1 6, 1 @, vii.4.49, H. A. Hunt (DAV). Madeline,
1, vii.12.25 (USNM). Poison L., 1 6, vi.13.41, C. W. Anderson (BERK).
Susanville, 1, viii.1.32, Blackwelder (USNM). Willow Ck., 1 6, 1 9, ix.9.58,
Allyn Smith (CAS). Los Angeles County. Claremont, 6 6,7 2°, Baker (CNG).
Lancaster, 3 m. N., 8, iv.3.53, R. K. Benjamin (USNM). Los Angeles, 5,
vii—.—, W. D. Richardson (USNM); 13, Coquillett (USNM); 1, ii—.19,
R. P. Dow (USNM). Palmdale, 1 2, vi.18.44, G. P. Mackenzie (LACM).
Pasadena, 2 6, 1 2, vi.27.16; 1 2, v.29.20 (AMNH). Pine Can., Hidden L.,
1 6,1 9, 11.15.53; 1 2, iii.21.53, Menke & Truxal (DAV). Pomona, 4, J. D.
Sherman, Jr. (USNM). San Franquito Can., 4 ¢, 11.29.53; 1 4, 1 9, iii.30.53;
24, 1 2, iv.1.53; 1 ¢, iv.3.53, L. A. Stange (DAV). Soledad Can., 1 ¢,
1.10.54, Menke & Sullivan (DAV). Tanbark Flat, 1 9, vii.13.50 (BERK).
Madera County. Chilkoot L., Jackass Mtns., 4 ¢, 2 9, vii.23-31.46, H. P.
Chandler (NMSU). Chowcilla, 18 m. E., 1 ¢, ii.2.64, H. B. Leech (CAS).
Coarsegold, 1 ¢, 1 2, H. P. Chandler (BERK). O’Neal’s, 1 4, iii.10.46, H. S.
Fitch (DAV). Windy Gap. 1, vii. 13.46, H. P. Chandler (CAS). Marin
County. Dillon Beach, 1 2, vi.24.63, H. B. Leech (CAS). Inverness, 2 6,
1 2, vii.20.50 (BERK). Pt. Reyes Sta., 2 ¢, x.1.60, J. S. Buckett (DAV).
Ross Valley, 1 2, ix.2.06, E. C. Van Dyke (AMNH). County, 2, J. D. Sher-
man, Jr. (USNM). Mariposa County. Miami Rngr. Sta., 1 2, vi.7.42, H. P.
Chandler (FM). Yosemite, 1 ¢, 1 2, vii.17.48, B. Adelson (BERK); 1 2,
v.24.64, W. W. Wade (CAS). Mendocino County. 1, v.31.17, E. R. Leach
(USNM). Albion, 1 m. S., 1 ¢, vii.1.63, H. B. Leech (CAS). Modoc County.
Cedarville, 5 m. E., 1 2, vi.5.60, J. Schuh (SCH). Juniper Flat, 1 6, vi.10.39,
J. J. Dubois (BERK). Mono County. Bodie, 8500 feet, 1 2, viii.13.62, H. B.
Leech (CAS). Bridgeport, Travertine Hot Springs, 2 m. S.E., 6700 feet, 1 ¢,
1 2, viii.11.62, H. B. Leech (CAS). Inyo Craters Lake, 8280 feet, 1 ¢,
vili.17.62, H. B. Leech (CAS). June Lake, 1 6, 5 2, ix.8-9.45, G. P. Mac-
kenzie (LACM). Leavitt Meadows, 7200 feet, 1 ¢, viii.10.62, H. B. Leech
(CAS). Mammoth (Lakes?), 4 ¢,5 @, ix.15.45, G. P. Mackenzie (LACM).
Twin Lakes, 1, vi.14.31, Blackwelder (USNM). Monterey County. Bryson,
2 2, iv.9.63, D. C. Rentz & K. A. Hale (CAS). Parkfield, 1 9, xi.20.63, H. B.
Leech (CAS). Salina(s?), R., 3, vii.17.25 (USNM). Napa County. Monti-
cello, Putah Ck., 2, xii.17.—, J. C. Bradley (CNL); 1 6, vii.22.30, A. T.
McClay (DAV); 2 8, 3 9, iii.30.31 (DAV). Nevada County. Hobart Mills,
1, H. D. Chandler (BERK). Sagehen Ck., 1 6, 1 2, x.15.55, J. L. Herring
(BERK). Webber L., 1 4, viii.22.64, E. S. Ball (SCH). Orange County.
Costa Mesa, 1 6, 2 2, iv.19.58, M. E. Irwin (DAV). Laguna Beach, 3,
vili.25-26.— (CNL); 1, vii—.21, C. T. Dodds (BERK). Laguna Can., 2 6,
xii.29.58, M. E. Irwin (DAV). Placer County. Tahoe, 14, vii—.— (USNM);
14, vii.6.09, J. D. Sherman, Jr. (USNM); 4 ¢6, 6 2, vii—— (AMNH).
Plumas County. Almanor Dam, 1 m. N.W., 2 6, 2 @, vii.17.47, C. H. Spitzer
(CAS). Buck’s Lake, 1 6, 1 9, vii.14.49, E. I. Schlinger (DAV). Chester,

JAMES R. ZIMMERMAN S1

1 9, vii.29.61, H. B. Leech (CAS). Clio, 1 2, viii.28.61, H. B. Leech (CAS).
Quincy, 4 m. W., 3 6, 4 8, vii.13.49, W. H. Wade (BERK); 1 ¢, 3 9, vii.6.49,
L. W. Isaak (DAV). Riverside County. Camp Pendleton, 2, x.7.45, H. P.
Chandler (BERK). Riverside, 3 6, 2 2, F. E. Winters (CAS); 6, vili.16.37,
Drake & Andre (USNM); 14, F. E. Winters (CNL). San Jacinto Mtns, 1 ¢,
2 2, —.—.32, F. E. Winters (CAS). San Jacinto Mtns., Keen Camp, 1 2,
R. F. Smith (FM). Sacramento County. Folsom, 1 2, iv.5.47, A. T. McClay
(DAV). Franklin, 1, iii.10.21, C. M. Packard (USNM). Sacramento, v.7.41,
A. T. McClay (DAV); 1 2, ii.—.20, H. F. Robinson (DAV). San Bernar-
dino County. Afton Canyon, 4, iv.18.31, Blackwelder (USNM). Barstow,
1 6 (NMSU); 2 6, 2 2 (AMNH); 7, viii.19.88 (ISU). Barton Flats, 2 9°,
x.4.58, M. E. Irwin (DAV). Colton, 1 (USNM). Cottonwood Sprgs., 7 6,
7 2, iv.5.48, MacSwain & Smith (BERK). Hesperia, 1 4, iv.7.39, G. P. Mac-
Kenzie (AMNH). Mtn. Pass, 1 ¢, iv.11.42, O. Bryant (CAS). Sweetwater
(Sprgs.?), 1, vi.2—, Dyar & Candell (USNM). San Benito County. Gon-
zales, 16 m. E., 2 9, iii.30.63, D. Reutz & K. Hale (CAS). San Diego County.
Anza-Borrego St. Pk., 1, iv—.53, A. & H. Dietrich (CNL). Banner, 6 m. E.,
1 6, vi.26.63, J. D. Birchim (CAS). Borrego (Sprgs.?), 1 6, iv.12.50, W. F.
Barr (FM). Campo, 1 2, iv.13.65, H. B. Leech (CAS). Descanso, 4 6, 1 2,
ix.23.52, 1, viii.7.14, J. C. Bradley (CNL). Escondido, 10 m. N., 5, xii.17.54;
12 m. N., 1, xi.20.53; 12 m. N., 1 2, iv.17.63, R. K. Benjamin (USNM).
Glenncliff Cp. (near Buckman Sprgs.), 3 6, 5 9, ix.21.54, Menke & Stange
(DAV). Laguna Mtns., near Buckman Sprgs., 2 6, 1 9, v.6.53, F. X. Wil-
liams (CAS). Mt. Palomar, 1 4, vi.21.59, M. E. Irwin (DAV); 2, x.21.45,
H. P. Chandler (BERK). Oceanside, Camp Pendleton, 1 2, x.7.45, H. P.
Chandler (FM). San Diego, 1, Mann (USNM); 1, G. H. Field (USNM); 2
(USNM); 2, iv.8.54, R. Straw (USNM). San Felipe Ck., 4 m. N., Scissors
Crossing, 2 2, iv.15.65, H. B. Leech (CAS). Sentenas Cn., 1 2, 1,26.56,
Menke & Strange (DAV). Warner’s (Springs), 1 2, viii.19.25 (MCZ). County,
1 9 (FM). San Mateo County. San Francisco Bay Marsh, Redwood City,
1 2, x.1.50, PSB (CAS). Woodside, Pulgas Temple, 4 ¢, 10 @, vi.10.51, PSB;
5 6,3 2, xi.4.51, PSB; 2 9, vi.23.52; 1 2, xi.25.54, PSB (CAS). Santa Bar-
bara County. Los Prietos, 4 m. E., 1 ¢, vi.26.65, M. R. Gardner (DAV).
Santa Barbara, 1, iv——, F. E. Winters (CNL): 4 6, 3 2, F. E. Winters
(CAS). Santa Clara County. Palo Alto, 2, ii.2.26, Blackwelder (USNM);
1 2, vi.6.92, V. L. Kellog (FM); 1, v.19.40, H. P. Chandler (BERK). Palo
Alto, Stanford U., 6 6, 1 2, ix.12.09, W. M. Mann (USNM). San Antonio
Val., 2, iii.2.48, J. W. MacSwain (BERK). San Jose, 1, Hubbard & Schwarz
(USNM). County, 1;2 ¢,2 2,C. F. Baker (USNM). Shasta County. Hat.
L., 1 6, vii.11.47, R. L. Usinger (BERK); 1 9, vii.11.47, E. E. Seibert (CAS);
Hat Ck., P.O., 1 2, vi.27.55, A.J. Mueller (DAV). Lake Britton, 1 ¢, vi.29.47,
C. H. Spitzer (CAS). Manzanita L., 5800 feet, 2 ¢, 3 2, x.1.44, B. Malkin
(FM); Manzanita L., Lassen Nt. Pk., 1 2, C. W. Anderson (BERK). Old
Station, 1 ¢, 1 9, vi.26.47, E. E. Seibert (BERK). Viola, 4 m. W., 2 4,
vi.27.47, E. E. Seibert (BERK). Sierra County. Sattley, 4 4, 3 2, vii.11.62,

MEM. AMER. ENT. SOC., 26

Sy THE GENUS LACCOPHILUS

R. L. Westcott (FM); 3 6,5 9, vii.11.62, Gill & Montgomery (DAV). Snag
Lake, 2 ¢, 3 Q, vii.5.64, J. E. Slansky (DAV). Siskiyou County. Grass Lake,
5000 feet, 1 6, vii.4.52, M. Cozier, W. Gertsch, R. Schrammel (AMNH).
Indian Tom L., 2 6, 2 9, ix.27.66, J. Schuh (SCH). Lower Klamath L., 4 9,
vi.1.57; 1 6, vi.6.66, J. Schuh (SCH). Montague, 10 m. S.E., 1 4, viii.27.64;
10 m. N.E., 1 9, viii.27.64, J. Schuh (SCH). Mt. Hebron, 5 m. S., Butte Ck.,
1 $, 3 2, v.1.66, J. Schuh (SCH). Weed, 4 m. S., 1 2, viii.12.58, J. Howell
(NMSU). Willow Ck., 1 2, vii.14.55, J. Schuh (MCHS); 1, ix.3.59, J. Schuh
(SCH); Willow Ck., Jewell Sprg., 1 9, ix.22.62, J. Schuh (SCH). Stanislaus
County. La Grange, 2.8 m. E., 2 ¢, 1 9, viii.14.62, H. B. Leech (CAS).
Sonoma County. Guerneville, 1, v.20.08, E. C. Van Dyke (USNM). Santa
Rosa, 2 (CNL). Solano County. Rio Vista, 1, iii.26.21, C. M. Packard
(USNM). Vacaville, 1 3, iv.15.43; 1 2, iv.8.49, A. T. McClay (DAV).
Trinity County. County, 1, ix.16.—, E. R. Leach (USNM); 1 6, viii.22.29,
E. R. Leech (CNG). Tulare County. Camp Potwisha, 4, vii.16.31, E. C.
Zimmerman (USNM). Kaweah, 4, Hopping (USNM). Porterville, 1 2,
Viii.19.62, E. E. Ball (SCH). Sequoia Ntl. Pk., 1 4, v.28.29; 1 9, v.19.30,
A. T. McClay (DAV). Three Rivers, 1, vii.12-14.07 (CNL). Tuolumne
County. Leland Mdw., 1 6, 1 9, viii.21.60, D. Q. Garagnaro (DAV). Near
Knights Ferry, 1 ¢, ix.22.63, H. B. Leech (CAS). Priest, 4.2 m. S.E., 1 6,
vili.19.62, H. B. Leech (CAS). Ventura County. Chuchupate Rgr. Sta., Fra-
zier Mtn., 1 2, v.2.59, C. W. O’Brien (BERK). Ojai, 1 4, vii—54, T. Barnes
(LACM). Yolo County. Davis, 1, iv—.49 (USNM); 1 6, ix.12.4, H. R.
Cameron (ORES); 1 9, iii.28.49, E. I. Schlinger; 1 2, iii.5.50, M. E. Taylor;
1 6, iv.18.59; 1 6, x.23.59, M. E. Irwin; 2 6, x.15.60; 1 2, ix.24.60, D. Q.
Caragnaro; 6 4, vi.26.61, V. L. Vesterby (DAV). Putah Cn., 1 2, iv.15.52,
A. A. Grigarick (DAV). COLORADO. Alamosa County. 1 4, vi.14.30;1 4,
1 2, vii.2.30, S. C. Bishop (NMSU); 1 4, iii.7—, F. C. Bowditch (MCZ).
Hooper, 9-10 m. E., San Luis Lakes, 2 6, 4 9, viii.13.65, H. B. Leech (CAS).
Archuleta County. Piedra, 3 4, vii.19.52, B. Malkin (FM). Chaffee County.
Buena Vista, 1, vii.6.96, Wickham (USNM). Conejos County. Conejos R. at
Hwy. 17, 1 ¢, 1 9, viii.12.65, H. B. Leech (CAS). N. Fork, Los Pinos R.,
at Hwy. 17, 1 6, viii.12.65, H. B. Leech (CAS). Denver County. Denver, 1,
vi.25.— (CNL); 10, x.—.35, A. Thrupp (CAS); 4 6, 2 2, vii.19.— (MCZ).
Garfield County. Glenwood Springs, 2 °, viii —— (MCZ). Gunnison County,
6, J. D. Sherman, Jr. (USNM). Gunnison, 1, ix.19.33, E. A. Andrews (CAS).
County, 7500 feet, 1 6, 1 2, vi.26-30.85, F. C. Bowditch (MCZ). Saguache
County. Mineral Hot Springs, 1 6, 2 9, viii.13.65, H. B. Leech (CAS). Sa-
guache, 24 ¢, 15 9, viii.18.52, B. Malkin & V.E.T. (FM); 10.5 m. N.W., 1 6,
vili.12.65, H. B. Leech; 12.5 m. N.W., 1 4, viii.12.65, H. B. Leech (CAS).
Weld County. Greeley, 1, Wickham (USNM). Adams County. Mesa, 2 m.
N. (Hwy. 95), 1 92, vii.20.6?, R. D. Anderson (RDA). Bear Lake County.
Paris (near), 1, vii.10.35 (CNG). IDAHO. Bingham County, 1, vii.24.—,
Hubbard & Schwarz (USNM). Blaine County. Clarendon Hot Springs, 5 é,
2 2, vii.22.52, B. Malkin (FM). Bonneville County. Idaho Falls, 1 ¢, 2 @
(CNG). Elmore County. Mountain Home, 1, vii.17.34, Lanchester (CAS).

JAMES R. ZIMMERMAN 53

Latah County. Moscow Mtn., 1, vii.20.38, H. M. Harris (ISU). Troy, 1,
vii.16.38, H. M. Harris (ISU). Nez Perce County. Lenore, 1000 feet, 1 6,
iv.1.51, W. F. Barr (FM). Oneida County. Malad City, 5 m. W., L. Malad R.,
1 6, 3 9, vii.13.52, B. Malkin (FM). Owyhee County. Hot Springs, 1 2,
vii.4.51, W. F. Barr (FM). Twin Falls County. Rogerson, 11 m. E., 2 9,
vii.20.52, B. Malkin (FM). MONTANA. Dawson County. Glendive, 1, Wick-
ham (USNM). Flathead County. Kalispell, 1, Wickham (USNM). Glacier
County. Cutbank, 2 6, 1 9, viii.22.54, P.S.B. (CAS). Jefferson County. 1 6,
vii.27.25 (USNM). Ravalli County. Corvallis, 2 2, v.20.52, A. A. Hubert
(USNM). Sanders County. Thompson Falls, 1 é, 1 2, vii.22-23.50, B. Mal-
kin (FM); 1, vii.22-23.50, B. Malkin (CAS). NEVADA. Elko County. Carlin,
2, vil.26.12 (USNM); 1, vii.17.12 (CNL). Lamoille Cn., 8 6, 24 9, vi.14.58,
F. D. Parker (DAV). Lincoln County. Hiko, Tonapah Jct., 1 6, vii.25.63,
P.S.B. (CAS). Nye County. Ash Meadows, 1 4, 1 2, iii.30.66, J. Schuh
(SCH). Beatty, 2m. N., 1 4, iii.29.66; 3 m. N. Amargosa R., 3 @, iii.29.66,
J. Schuh (SCH). Warm Springs, 6 6, 5 9, vii.25.63, P.S.B. (CAS). Ormsby
County. Carson City, 3, vii.30.—, Wickham (USNM). Washoe County.
IXL Ranch, 2 9, vii—.51, N. Pasquale (ORES). Reno, 1 6, 1 2 (AMNH);
2 2, 1.2.57, F. Parker (DAV); 1 6, Wickham (ANSP); 1, Wickham (USNM).
OREGON. Benton County. Buchanon, 4 6,5 2, ix.10.49, V. Roth (ORES).
Corvallis, 1 2, vili.13.35; 1 9, x.18.35, G. Ferguson (ORES); 3 2, xi.2.06
(ORES); 1 2, x.28.06, Wood (ORES); 2 6, 1 9, xi.1.08, Hays, Sprague &
Reynolds (ORES); 3 9, vi.23.—; vii.4.09, iv.5.36 (ORES); 1 2, iii.30.30, R. L.
West (ORES); 2 6, 3 2, x.24.—, xi.3.33, J. Schuh (ORES); 1 4, v.26.35,
I. Tarshis (ORES); 2 6, 2 9, iii.7-16.36, N. D. Larsen (ORES); 1 2, iv.10.38,
S. Jewett (ORES). Corvallis, Oak Ck., 1 6,1 9, v—.39, G. G. Black (ORES).
Corvallis, Colorado L., 4 9, iv.7.30, L. Seghetti (ORES). 3, iv.22.09, G. F.
Moznette (CNL); 3 (USNM); 5, iv.25.—, J. D. Sherman, Jr. (USNM); 44,
J. D. Sherman, Jr. (USNM); 21, 1, iv.28.01, J. D. Sherman, Jr. (USNM); 8,
Vita O09 sD» Shermans in (USNIM) lid.) 2498 iv.27-— 146. 119)
vi.24.—; 10 6, 1 9, vii.18—; 4 ¢, 3 9, ix.20.—; A. R. Woodcock (AMNH).
Summit Prairie, 4 ¢, 6 9, viii.9.39, Gray & Schuh (ORES). Yew CKk., near
Mt. Alsea, 2 ¢, v.25.45, G. L. Bennett (ORES). Coos County. Coos Bay,
1 ¢, 1 9, vii.9-10.51, B. Malkin (FM). Curry County. Pistol R., 6 6, 6 2,
ix.18.50; 1 6, vii.7.51, B. Malkin (FM); 1, vii.7.51 (CAS). Deschutes County.
Bend, 4, vii.23.32, R. Blackwelder (USNM). Bend, 20 m. S., 2 4,7 9, iii.8.39,
Gray & Schuh (ORES). Near Tumalo, Deschutes R., 3 4, 3 2, iii.22.39, Gray
& Schuh (ORES). Douglas County. Gardiner, 8 m. E., Otter Slough, 2 @,
vi.1.66, J. Schuh & J. Vertrees (SCH). Grant County. Kimberly, 15 m. S.,
2 6, vii.1.60, J. D. Lattin (ORES). Silvies Valley, 2 4, iv.28.57, J. Schuh
(SCH). Harney County. Blitzen Valley, 1 4, 4 9, viii.1.36, S. Jewett (ORES).
Buena Vista, 1 6, v.19.56, J. D. Lattin (ORES). Burns, 25 m. E., 2 2,
vili.21.45, H. A. Scullen (ORES). Burns, 15 m. S., 1 &, vii.13.53, Roth &
Beer (ORES). Burns, 3 6, 5 ?, vi.21.51, B. Malkin (FM). Malheur, L.,
1 ¢, vi.20-21.51, B. Malkin (FM). Malheur Wldlf. Ref., 1 6, viii.15.62

MEM. AMER. ENT. SOC., 26

54 THE GENUS LACCOPHILUS

(MCHS). Jackson County. Trail, 1 ¢, 292, 1x.9.50, Malkin & Thatcher
(FM); 1, ix.9.50 (CAS). Medford, 3 6, 3 9, iv.1.39, F. Lawrence (DAV).
Josephine County. Grants Pass, 1 6, vii.3.06, Eismann (ORES). Rogue River,
1, 1x.8.02, Biederman (USNM). County, 1 2, vi.6.34, F. W. Nunenmacher
(FM). Klamath County. Aspen L., 3 ¢, 3 2, v.5.57, J. Smith (SCH). Bly,
5 m. E., 4 6, v.5.62, R. Scott & J. Schuh. Bly Mt., 2 6, 4 9, ix.15.60, J.
Schuh (SCH). Chiloquin, 12 m. E., Sprague R., 1 6, 3 9, vii.1-3.51, B. Mal-
kin (FM). Chiloquin, 7 m. N., Williamson R., 1 ¢, 2 9, ix.23.66; 6 m. E.,
1 ¢, 1 9, ix.21.65, J. Schuh (SCH). Crescent, 4 ¢, 5 2, ix.10.50, B. Malkin
(FM); 1 9, v.12.57; 1 2, J. Schuh (SCH); 2 m. N.E., 2 2, v.13.66; 11 m.
N.E., 1 2, v.13.66, J. Schuh; 13 m. S.E. Bly, 1 2, iv.26.62, Vertrees & Schuh
(SCH). Ft. Klamath, 5 ¢, 3 9, viii.10.50, B. Malkin (FM). Klamath Falls,
10 m. N.W., 1 6, vi.16.52 (ORES); 1 6, iv.22.57; 1 6, v.2.57; 1 6, iv.23.53;
1 2, v.31.48, J. Schuh (ORES); Modoc Pt., 1 2, vi.1.55, Schuh (ORES); Poe
Valley, 1 9, v.27.55, Schuh (ORES); 5 6, 2 2, ix.3.50, B. Malkin (FM).
Klamath (?), 1, G. M. Greene (USNM). Mare’s Egg Spring, 3 4, ix.7.59,
J. Schuh (SCH). Méidlland, 3 m. S., 20 6, 13 92, ix.3.50, B. Malkin (FM).
Poe Valley, 4 6, v.13.66, J. Schuh (SCH). Lake County. Abert L., 1 ¢,3 9°,
vi.7.58, Vertrees & Schuh (SCH). Chandler St. Pk., 4 6, vi.29-30.51, B. Mal-
kin (FM). Near Valley Fall, Chewaucan R., 1 6, 1 9, vi.7.55, J. Schuh
(ORES); 1 ¢, 11 Q, iii.16.39, Schuh & Gray (ORES). Lakeview, 1 9, vi.27-
28.51, B. Malkin (FM); 1 4, vi.—.58, J. Schuh (SCH). Silver L., 10 m. N.,
3 2, v.12.57, J. Schuh (SCH). Summer L., 1 2, v.18.57, J. D. Lattin (ORES).
Lane County. Eugene, 2 2, vi.26.41; 2 @, viii.2.41; 1 6, 1 2, iv.5.42; 2 9,
iv.11.42; 2 4, vii.13-14.51, B. Malkin (FM). Linn County. Albany, 1,
vi.15-28.—, Wickham (USNM). Tangent, 2 ¢, 3 2, v.4.38, Edman (ORES).
Malheur County. Drewsey, 4 m. E., 1 6, 1 @, ix.10.64, J. Schuh & W. Peters
(SCH). Multnomah County. Gaston, 4, J. D. Sherman, Jr. (USNM). Port-
land, 1, J. D. Sherman, Jr. (USNM). Wasco County. Simnasho Hot Springs,
14 m. S., 1 4, v.22.59, J. D. Lattin (ORES). Washington County. Forest
Grove, 1 6, x.—.06, J. C. Birdwell (ORES). Yamhill County. McMinnville,
1 4, vii.—.36, K. Fender (FM); 1 ¢, v.—.38, K. M. & B. M. Fender (LACM).
TEXAS. Hale County. Plainview, 1 6, vii.14.54, F. C. Harmston (RDA).
UTAH. Boxelder County. Rabbit Sp. Reservoir, 1 6, 1 2, vii.2.59, R. D.
Anderson (RDA). Showville, 1 6, 1 @, viii.1.59, R. D. Anderson (RDA).
Cache County. Benson, Logan Meadows, 13, viii.7.52, Knowlton & Wood
(USNM). Wasatch Range, Beaver Mtn., 7700 feet, 1 4, vil.6.52, B. Malkin
(FM). Juab County. Callao, 1 6, 1 2, xii.27.57, R. D. Anderson (RDA).
Sevier R., at Hwy. 91, 1 6, 1 9, vi.11.59, G. Musser (RDA). Kane County.
Zion Ntl. Pk., 2 m. E., 2 2, vi.21.59, R. D. Anderson (RDA). Piute County.
Greenwich, 5 m. S., 1 2, viii.3.64, H. B. Leech (CAS). Rich County. Bear
L., 1 ¢, vii.4.59, R. D. Anderson (RDA). Salt Lake County. Saltair, 1,
vi.15.41, H. P. Chandler (CAS). Salt Lake, 1, iv.7.—; vi.13.—, Hubbard &
Schwarz (USNM). Salt Lake City, 1 9, vii.14.52, B. Malkin & S. Mulaik
(FM). Salt Lake City, Jordan R., 3 6, 1 9, iii.19.58; 2 $, 1 9, iv.20.57,
R. D. Anderson (RDA). Summit County. Kimball Jct., 3 m. W., 1 2, v.12.57,

JAMES R. ZIMMERMAN 55

R. D. Anderson (RDA). Pineview, 1 m. E., 1 2, vii.2.59, R. D. Anderson
(RDA). Tooele County. Mills, 3 m. S., 2 6, 3 2, v.2.58, R. D. Anderson
(RDA). Unita County. County, 2 2, G. E. Wallace (CNG). Utah County.
Goshen, 1 m. W., 2 4, iv.27.58, R. D. Anderson (RDA). Provo, 6, x.10.36,
H. P. Chandler (CAS). Utah L., 13, vi.10.—, Hubbard & Schwarz (USNM);
1 6, vi.l9— (ANSP). Utah L., American Fork, 1 2, iv.23.58, R. D. Ander-
son (RDA). Washington County. Hurricane, 1 2, vi.22.59, R. D. Anderson
(RDA). Near Central, Santa Clara Ck., 2 6, 2 2, v.10.58, R. D. Anderson
(RDA). New Harmony, 1 ¢, 3 9, x.11.64, R. D. Anderson (RDA). St.
George, 1, vii.—.—, Wickham (USNM); 12, A. M. Woodbury (USNM); 2 ¢,
5 2, iv.—.25 (USNM); 1 m. W., 3 6, 3 9, xii.29.59, G. Musser (RDA).
Virgin R., 1 6,2 2, G. Weldt (AMNH). Weber County. Ogden, 8, vii.3.—,
Hubbard & Schwarz (USNM). WASHINGTON. Adams County. Ritzville,
4, vili.29.24, D. Dunavan (USNM); 2, vili.10.21, M. C. Lane (CAS). Benton
County. Pasco, 4 6, iii.26.60, J. Shepard (ORES). Chelan County. Leaven-
worth, 8, v.29.42, J. G. Needham (CNL). Douglas County. Stratford, 2,
ix.3.20, M. C. Lane (USNM). Grant County. Vantage, E. Side Columbia R.,
2 2, vii.—.53, Crabtree (FM). Jefferson County. Pt. Townsend, 1 ¢, 1 2
(AMNH). King County. Issoquady, 1 ¢, ii.5.41, H. J. Jensen (BERK).
Seattle, 1, iv.3.72 (CNL). Skykomish R., 1, vi.1.92 (USNM). Seattle, 1 2,
iv.19.45, H. J. Jensen (BERK); 1 2, v.12.— (ANSP). Kitsap County. Wins-
low, 1, ix.10.—, Wickham (USNM). Kittitas County. Ellensburg, 1 2, 1x.26.54,
B. Malkin (FM). Vantage, 8 m. N., Whiskey Dick Can., 1 2, viii.4.54, B.
Malkin (FM). Okonagan County. Okonagan, 4, iv.25.29, M. D. Leonard
(CNL). Pierce County. Buckley, 1, v.28.42, J. G. Needham (CNL). Ft.
Lewis, 24, vii.7.51, R. Schuster (BERK). Snohomish County. Chase Lake,
T G5 UV Qo WWULSI4E D Go Il Oy i HS98 il Sy il @5 wes I Gs, 1) 5 xail/es
B. Malkin (FM). Sultan, 1 6, 1 92, vi.1.53, B. Malkin & D. Boddy (FM).
Walla Walla County. College Place, 1 6, v.2.45, W. Blehm (MCHS). Walla
Walla, 1 6, ix.2.26, M. W. Stone (ORES). Whitman County. Pullman, 3,
vili.17.01, C. Y. Piper (USNM); 1, vi.20.40, C. Yocum (ISU). Yakima
County. Toppenish, 1, iii.20.25; 2, vii.20-27.24, D. Dunavan (USNM). Wash-
ington County. Terry, 50, Morrison (USNM). East Washington, 2, C. V.
Piper (USNM). WYOMING. Albany County. Rock Creek, 1, viii.24.41, H. P.
Chandler (CAS). Converse County. Douglas, 4.5 m. W., 1 6, 1 2, vii.27.64,
H. B. Leech (CAS). Crook County. Devils Tower N. M., 4m. S., 5 6, 6 2,
vill.4.52, B. Malkin (FM). Goshen County. Ft. Laramie, Laramie R., 1 ¢,
4 2, vii.29.64, H. B. Leech (CAS). Ft. Laramie, Natl. His. Site, 1 ¢, 1 2,
vili.19.65, H. B. Leech (CAS). Ft. Laramie, 5.3 m. E., 3 2, viii.18.65, H. B.
Leech; Jay Jay Em, 2.5 m. N., 3 @, viii.20.65, H. B. Leech (CAS). Lingle,
3.8 m. E., 1 6, viii.19.65, H. B. Leech (CAS). Johnson County. Crazy
Woman Ck., S. Fork, 1 6,2 2, vii.26.64, H. B. Leech (CAS). Laramie County.
Cheyenne, 12.5 m. N.E., Lodgepole Ck., 3 6, 2 2, viii.18.65, H. B. Leech
(CAS). Natrona County. Natrona, 2.5 m. S.E., 1 2, viii.20.65, H. B. Leech
(CAS). Midwest, 8.5 m. N.W., Duquot Ck., 1 6, 2 2, vii.27.64, H. B.

MEM. AMER. ENT. SOC.. 26

56 THE GENUS LACCOPHILUS

Leech (CAS). Niobrara County. Manville, 3 m. W., 2 ¢, viii.20.65, H. B.
Leech (CAS). Platte County. Glendo, 2 m. S., Horseshoe Ck., 1 é, 2 9,
vii.29.64, H. B. Leech (CAS). Sweetwater County. Farson, 1 6, viii.22.65,
H. B. Leech (CAS). Green River, | ¢, 1933, Wickham (USNM). Rock
Springs, 10 m. W., 1 ¢, vili.23.65, H. B. Leech (CAS).

MEXICO. — BAJA CALIFORNIA. La Suerte, 2 6, vi.4.63, R. K. Benjamin
(CAS); on southern slope of Sierra San Pedro Martir, Hamilton Ranch, 8 m.
upstream near Colonia Guerro, 1 92, iv.23.63, H. B. Leech (CAS). San Felipe,
1, J. R. Sherman, Jr. (USNM). Tijuana, 45 km. S., 1 ¢, 1 9, vili.21.53, L. A.
Stange (DAV).

Laccophilus maculosus shermani Leech, new status
(Figs: 355365276520)

Laccophilus shermani Leech, 1944, pp. 4-6. Holotype: male; Bear Canyon,
Santa Catalina Mountains, Pima County, Arizona; iv.2.38, E. C. Van Dyke,
Canadian National Collection, Ottawa.

DIAGNOSIS. — L. m. shermani belongs to the irrorated species with saw-
like ovipositors and metacoxal files. The venter is yellow to reddish-brown, and
the elytral pattern is relatively uniformly irrorate except for maculae. L. mis-
tecus aztecus, which occurs in central Mexico, has much the same color and
pattern, but differs in aedeagus, its larger size (0.2 mm longer), and lack of
female epipleural flange. L. fasciatus subspecies are smaller (see diagnosis for
decipiens) and have black markings on the posterior half of the elytra. L. fusci-
pennis is evenly irrorate and characteristically has a flange, but lacks clear
maculae on the elytra. It is about one millimeter shorter (average length, 4.5
mm). L. proximus is about the same length or even shorter than L. fuscipennis.

NOMENCLATURAL NOTES. — No synonymical confusion ex-
ists for this race. It was considered to be part of decipiens, until
Leech recognized that the male aedeagus and coxal file were differ-
ent. Mexican records other than Baja California undoubtedly refer
to shermani.

VARIATION. — The mean length of males is about 5.8 mm and
that of females is about 5.6 mm — about the same as for decipiens.
The most southeastern sample (Puebla) has high mean values for all
measurements, indicating that there could be a change in size in that
peripheral population; but since there are only seven specimens of
each sex available, it is not possible to give a conclusive answer. The
nearest sample (Huasca, Hidalgo) gives values comparable to the rest
of the race. The PW/EL ratio shows no geographical trend. In
general, shermani presents much the same picture as decipiens and
maculosus; i.e., widespread, outbreeding populations with little re-
gional specialization within the race.

JAMES R. ZIMMERMAN 57

The female epipleural flange is the most striking variable char-
acter in shermani, but the only obvious geographical relationship is
that it decreases near the northern limit of the race. This decrease
suggests influence from decipiens.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— The range of L. m. shermani covers Arizona, New Mexico, west
Texas, and the Mexican Plateau. It is limited on the northeast by
the High Plains of New Mexico and on the northwest by the lower
Colorado River Valley. In Mexico, it is almost entirely confined to
the Central Plateau; but I have collected it once near Mazatlan.
Numerous collections in the lowlands of Sonora have not produced
any shermani, however. It occurs near Monterrey and Saltillo in the
Sierra Madre Oriental, but not in the lowlands to the east. The
range’s southernmost extensions are in Puebla and Michoacan.

L. m. shermani is most commonly found in the Pinon-Juniper
zone or lower Ponderosa zone of New Mexico, Arizona, and West
Texas; but it does occur in the flats between mountains. It appears
to have less ecological tolerance than decipiens. In Mexico, it is most
frequently found in pine-oak associations or in the pools of streams
not exceeding 8000 feet.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Chiraca-
hua Mtns., N. Portal, 1 ¢, 1 2, viii.13.52, H. B. Leech (CAS). Chiracahua
Mtns., Herb Martyr L., 3 ¢, 8 9, ix.5.61; Cave Canyon Ck., 2 6, 3 9, 1x.5.61,
JRZ (NMSU). Douglas, 1 2, iii.15.33, W. W. Jones (BERK); 13 m. E., 1 2,
vi.11.59, L. A. Stange (DAV); 6m. E., 1 6, 1 2, v.18.63, K. L. McWilliams
(NMSU). Fairbanks, 2, ili.12.36, J. G. Needham (CNL). Huachucha Mtns.,
Sunnyside Cany., 1 6, 2 9, viii.4.52, Leech & Green (CAS). Huachucha City,
14 6,8 9, ix.6.61, JRZ (NMSU). Palmerlee, 2, viii—— (USNM). Portal,
SWRS, 5400 feet, 5 m. W., 1 2, iv.23.56, E. Ordway (AMNH); Portal, 2 9,
vii.20.58, J. M. Marston (DAV); Portal, 5 m. W., 1 6, 1 2, viii.17.58, J. M.
Marston (DAV). St. David, 1 2, vii.17.45, G. E. Pickford (FM); 7 6, 12 2,
vii.25.61, JRZ (NMSU). Tombstone, 2 ¢, vii.7.44, G. E. Pickford (FM).
Coconino County. Flagstaff, 7 m. S., 3 6, 3 2, iv.7.66, J. Schuh (SCH).
Grand Canyon, Supai, 3500 feet, 8 4, 20 9, viii.2-4.34, E. L. Bell; Phantom
Ranch, 2500 feet, 1 ¢,4 9, vii.26.34, D. Rockerfeller (AMNH). Williams, 3,
i1.6.—, Barber & Schwarz (USNM). Gila County. Carrizo Ck., 23 6, 31 2,
vii.20.63, PSB (CAS). Globe, Pinal Ck., 4, iv.25.53, A. & H. Dietrich (CNL);
2 2, v.10-11.25, D. K. Duncan (CAS). Graham County. Galiuro Mtns., 1,

MEM. AMER. ENT. SOC., 26

58 THE GENUS LACCOPHILUS

yv.21.—, Hubbard & Schwarz (USNM). Geronimo, 2 9°, iv.28.24, J. O. Martin
(CAS). Graham Mtns., 9500 feet, Snow Flat, 5 6, 1 2, ix.13.52, B. Malkin
(FM). Graham Mtns., Hwy. 666 at Marijilda Canyon, 2 6, 8 Q, viii.3.65,
H. B. Leech (CAS). Safford, 10 m. S., 1 é, ii.2.36, O. Bryant (CAS). Green-
lee County. Hardy, 1, Wickham (USNM). Maricopa County. Phoenix, 1
(USNM); 34 6, 24 2 (AMNH). Wickenburg, 5 m. E., Hassayampa R., 2 2,
iv.2.66, J. Schuh (SCH). Mohave County. Peach Springs, 2, —.—.33; 1,
vili.27.—, Wickham (USNM). Navajo County. Clay Springs, 35 6, 37 2,
vii.21.63, PSB (CAS). Lakeside, 2, vi.11.46, G. P. Mackenzie (CAS); 1 6,
3 2, vill.22.52, B. Malkin (FM). Pima County. Arivaca, Arivaca Ck., 1 9,
vul.31.52, J. W. Greene (CAS). Lowell Rngr. Sta., 2700 feet, 5 ¢, 7 2,
vii.6-20.16 (AMNH). San Fernando (now Sasabe), 3, vili.28.25, R. Budlong
(USNM). San Xavier Mission, 1, D. K. Duncan (USNM— paratype). Tucson,
1 2, 1ix.11.24, A. A. Nichol (CAS); 1 6, vi.1.33; 2 6.2 2, ix.—.33, O. Bryant
(CAS); 1 6, 3 2, 1.18.46, J. W. Green (DAV); 3, Shoemaker (USNM); 1,
xi.—.24; 1, x1.4.24, Frost; 1, xi.8.27, L. C. Bailey; 1, xi.8.34, K. Thomas;
1, v.22.32 (ARI). Tucson, Santa Catalina Mtns., 4, xi.15.24; 1, x.21.24; 2,
xil.4.24, A. A. Nichol (ARI). 1, ix.17.57, C. W. Nichol & O’Brien (ARI);
4, iv.S-18.—, Hubbard & Schwarz (USNM). Tucson, Santa Catalina Mtns.,
Bear Cany., 1 6, xi.15.24, A. A. Nichol; 3 ¢, 8 9, 1.2.38, Van Dyke (CAS);
1, x1.4.34, L. P. Wehrle; 1, xi.5.35, H. H. Cole; 2, x.16.43, J. Hendrickson
(ARI); Tucson, Santa Catalina Mtns., Sabino Canyon, 1, iii.22.52, R. L. Catsitt
(ARI); 1, J. W. Tilden (USNM — paratype). Tucson, Santa Catalina Mtns.,
Molina Basin, 1 2, vi.l1.58, MacNeil & MacNeil. Tucson, Santa Catalina
Mtns., Lowell Rang. Sta., 8, vii.6-20.16 (USNM). County, 1 6, x.21.24,
A. A. Nichol (BERK). Quitobaquito, O.P.C., Natl. Mon., 2, iv.25.53; 5,
v.1.53, A. & H. Dietrich (CNL). Pinal County. Aravaipa Cn., near Mam-
moth, 1, v.21.53, A. & H. Dietrich (CNL). Santa Cruz County. Atasco Mtn.,
1, 1x.26.53, G. D. Butler (ARI). Tumacacori, Sycamore Cn., 1 6, 4 2, viii.3.52,
H. B. Leech (CAS). Near Ruby, 1, xi.20.55, G. D. Butler & F. G. Werner
(ARI). Madera Cn., 1 6, vii.3.63, V. L. Vesterby (DAV). Nogales, 1 é,
2 2, vuli.19.06; ix.30.06, Blaisdell (CAS); 1 2, viii.19.06, F. W. Nunenmacher
(FM). Pajarito Mtns., Pena Blanca, 33 6, 26 2, vii.19.-vilil.8.62, R. Arnett
& E. Van Tassel (CUA). Pena Blanca Lk., 2 6,3 92, iv.4.66, J. Schuh (SCH).
Patagonia, 4m. S., 1 ¢, 3 9, iv.4.66, J. Schuh (SCH). Santa Rita Mtns., Box
Can., 2 6, viii.29.52, B. Malkin (FM); Santa Rita Mtns., Florida Canyon,
4 8,1 9, 1.2.62; Santa Rita Mtns., 1, v.29.—, Hubbard & Schwarz (USNM).
Tumacacori, Sycamore Cn., 2, vi.3.53, A. & H. Dietrich (CNL). Yavapai
County. Bloody Basin, 1 @, vi.8.47, F. H. Parker (CAS). Castle Hot Springs,
1 6, iv.5S.42 (AMNH). Minehaha, 1 (USNM); 12 (AMNH). Prescott, 3,
Vill—.— (USNM); 17 6, 26 9, viii ——- (AMNH). NEW MEXICO. Berna-
lillo County. Albuquerque, 2, C. V. Riley (USNM); 2, M. L. Linell (USNM);
1, vii.2.—, Wickham (USNM); 1 6, 2 2; 14, 19, Wickham (AMNH); 3,
vii.7.88 (ISU). Catron County. Alma, 15 m. N., 3 6, 5 9, vii.16.63, R. D.
Ohmart (NMSU). Datil, 8 m. S., 4 6, 4 2, vii.17.63, R. D. Ohmart (NMSU).
Quemado, | 4, ix.1.41, O. Bryant (CAS); 16 46, 16 2, vii.16.63, R. D. Ohmart

JAMES R. ZIMMERMAN 59

(NMSU). Colfax County. Near Maxwell, 5 6, vii.l4.61, JRZ (NMSU).
Raton, 1 ¢, iv.25.51, O. Bryant (CAS); 23 m. S.E., 2 6, vii.5.63, R. N. Gen-
naro (NMSU). Van Houten, 7000 feet, 5 ¢, 5 2, vii.6.63, R. N. Gennaro
(NMSU). Dona Ana County. Las Cruces, 1 ¢, vil.9.61, JIRZ (NMSU); 1 ¢,
vii.6.61, E. Staffeldt & JRZ (NMSU); 1 2, x.1.62, K. L. McWilliams (NMSU).
Mesilla, 1 46, vii.6.61, JRZ (NMSU). Grant County. Silver City, 10 m. W.,
33 6, 50 2, vil.16.63, R. D. Ohmart (NMSU). Harding County. Gallegos,
41 m. N., 1 2, x.23.65, A. H. Smith & T. O. Boswell (NMSU). Hidalgo
County. Animas, 12 m. S.E., 1 6,2 2, K. L. McWilliams (NMSU). Animas
Mtns., Double Adobe Ranch (near Animas), 1 ¢é, 1 Q, vili.15.52; 6 4, 4 Q,
vii.31.65, H. B. Leech (CAS). Lincoln County. Alto (north of Ruidoso),
13 6, 12 Q, vu.27.61, JRZ (NMSU). Angus, Rio Bonito, 1 6, 4 2, vili.7.65,
H. B. Leech (CAS). Luna County. Deming, 15 m. N.W., Mimbres R., 9 é,
4 2, vil.17.61, JRZ; Deming, 10 m. W., Hwy. 26, 1 2, iv.20.63, K. L. McWil-
liams; Hurley, 4 ¢, 1 2, vii.17.61, JRZ; Lordsburg, 13 m. E., 1 92, K. L.
McWilliams (NMSU). McKinley County. Ft. Wingate, 26 (USNM). San-
doval County. Cuba, 3 m. S.W., 5 6, 7 Q, vii.11.63, R. D. Ohmart; Jemez,
Jemez Ck., 1 2, iv.30.57, A. Smith (NMSU). Sierra County. Arrey, 7.5 m.
W., 14 6, 4 2, 11.20.63, K. L. McWilliams (NMSU). Hillsboro, 4 m. E.
(label says Pedra R. Gorge in Mimbres Mtns., but must mean Percha Ck.),
3.6, 1 9, viii.31.52, B. Malkin (FM); 1 ¢, iv.7.63; Hillsboro, 1 ¢, iv.7.63,
JRZ (NMSU). Socorro County. Near Bingham, 1 ¢, 3 Q, vii.21.62, R. D.
Ohmart (NMSU). Torrance County. Mountainair, 10 m. S., Abo State Mon.,
1 9, vili.21.67, H. B. Leech (CAS). Valencia County. Bluewater, 1, 1933,
Wickham (USNM). Fence L., 4 m.S., 3 2, vil.18.63, R. D. Ohmart (NMSU).
TEXAS. Brewster County. Marathon, 4 m. S., 1 ¢, ix.5.52, B. Malkin (FM).
Culberson County. Nickel Creek Sta., 2 m. E., 11 6, 8 2, ix.2.52, B. Malkin;
1 6, 1x.2.52, B. Malkin & V. Thatcher; 1 2, ix.9.52, B. Malkin (FM). El Paso
County. Clint, 1 m. W., 11 6, 5 2, vii.19.61, JRZ (NMSU). Jeff Davis
County. Davis Mtns., 1 (CNL); 1, vii.16.41, B. E. White (CNL); 1 4, 1 2,
v.9.51;2 6, 1 2, x.11.49, O. Bryant (CAS); Davis Mtns., Limpia Can., 14 4,
12 9, 1v.19.53, B. L. Adelson & M. Wasbauer (BERK); 11 6, 10 2, iv.19.53,
A. J. Adelson & M. Wasbauer (CAS); 1 ¢, ix.4-5.52; Davis, 13 m. S., Kent,
4 6,1 9, ix.8.52; 1 ¢, ix.13.52, B. Malkin (FM). Davis Mtns., Elbow Can.
Ck., 10 6, 9 2, x.27.61; Madera Can. Ck., 2 6, viii.3.61; Limpia Can. Ck., 3 6,
3 2, vili.3.61, JRZ (NMSU). Ft. Davis, 1, vi.10.14, C. Thompson (USNM);
2 6,3 Q, viii.3.61 (NMSU).

MEXICO.— AGUASCALIENTES. Aguascalientes, 1 m. N.,5 6,5 9, vii.26.62,
JRZ (NMSU). CHIHUAHUA. Chihuahua City, 1 ¢, Hége (AMNH) (ex.
B. C. A. Coll.) ; Chihuahua, 43 m. N., 18 6, 19 2, xii.8.62; Camargo, 1 m. N.,
1 6, vii.25.62; Casas Grande, 5 m. E., 9 6, 11 92, vii.20.64; Colonia Juarez,
2m.S.W., 11 6, 16 2, xii.20.64; Hidalgo de Parral, 2 m. S., 1 6, 2 2 vii.25.62;
10 m. S., 1 6, 2 9, xii.9.62; JRZ (NMSU). Primavera, 5 ¢, 4 2, vi.30.47,
M. Cazier, R. Shrammel, W. Gertsch (AMNH). Santa Barbara, 63 m. W.,
5500 feet, 1 3, vii.20.47, Gertsch (AMNH). Zaragosa, 1 6, 1 9, vii.19.61,

MEM. AMER. ENT. SOC., 26

60 THE GENUS LACCOPHILUS

JRZ (NMSU). COAHUILA. Ramos Arizpe, 5 6,7 2, vii.7.63, JRZ (NMSU).
DURANGO, 1, Wickham (USNM). Durango, 20 m. W., 4 6,7 9, xii.10.62,
JRZ (NMSU). EI Pino, 5m. N., 1 6,1 2, J. D. Lattin (BERK). LaZarca,
15 m.N., 15 ¢, 17 92, vii.25.62; Llano Grande, 10 m. E., 1 ¢, 1 9, xii.10.62;
Nombre de Dios, 1 ¢, 4 2, vii.26.62; San Juan Del Rio, 10 m. N., 2 6, 3 9,
xii.9.62, JRZ (NMSU). GUANAJUATO. Irapuato, 5 m. S., 1 2, vii.27.62,
JRZ (CAS). HIDALGO. Near Huasca (San Miguel Regla), 3 6, 1 2, vii.10.63;
Zimapan, 1 6, 2 2, vii.9.63, JRZ (NMSU). MICHOACAN. _§Insurgentes
Morelos Pk., 1 2, vii.27.62, JRZ (NMSU). NUEVO LEON. Sta. Catarina,
Huasteca Can., 3 6, 4 9, vii.7.63, JRZ (NMSU). PUEBLA. Acatzingo, 2 ¢,
1 Q, vili.28.62; Cuapiaxtla 5 6, 6 9, viii.24.62, JRZ. San Martin Texmelu-
can, 1 6,1 @, viii.24.62, JRZ (NMSU). QUERETARO. San Juan Del Rio,
1 6, 2 9, iii.27.63, JIRZ (NMSU). SAN LUIS POTOSI. San Luis Potosi,
2m.S., 1 4, 3 2, xi.21.48, H. B. Leech (CAS). Santa Maria del Rio, 2 ¢,
3 2, ili.26.63, JRZ (NMSU). SINALOA. Mazatlan, 7 m. S., 4 6, 4 2,
xii.11.62, JIRZ (NMSU). SONORA. Bavispe, 1 4, vii.26.63, JRZ (NMSU).
ZACATECAS. Fresnillo, 61 m. S., 1 ¢, 1 2, vi.25.54, R. H. Brewer (CAS);
16 m. N.W., Rio Trujillo, 1 ¢, 1 2, vi.29.54 (CAS); 45 m. W., 7700 feet,
14, 2 9, vi.25.54, R. H. Brewer (CAS). Laguna Balderama, 8200 feet,
vii.8.54, R. H. Brewer (CAS). Pinos, 10 m. S., 1 2, viii.1.59, Ray Bandar
(CAS). Sombrerete, 10 m. W., 1 6, 4 9, vii.1.54, R. H. Brewer (CAS).

INTERGRADATION IN LACCOPHILUS MACULOSUS

There are two intergrade zones in Laccophilus maculosus — one
between maculosus and decipiens and the other between decipiens
and shermani. Intergrades between maculosus and decipiens occur
in a band from northeastern Colorado, through Nebraska and South
Dakota, and into southern Manitoba. Figure 280 is an illustration
of an intergrade elytral pattern from Colorado. Presumably, inter-
grades also occur in eastern Wyoming and western North Dakota;
but they have not yet been recognized. It is not surprising that they
have not been detected, however. Intergrades can be difficult to de-
termine in many instances, and even large samples usually include
only a few individuals that are intermediate for several characters.
Most populations seem to be composed mainly of F2’s or backcrosses
with one of the parental races.

Intergrades between decipiens and shermani occur in a narrow
zone extending from north central New Mexico through northern
Arizona to a single, indefinitely placed locality in eastern Riverside
County in California. These intergrades could be verified only if the
aedeagus was intermediate. The elytral patterns are too similar to
show intergradation. The female epipleural flange should be helpful

JAMES R. ZIMMERMAN 61

shermani

30 decipiens

N U M BER §S

maculosus

YE

4.8 5.0 5.2 ; 5.6 5.8 6.0 6.2
TOTAL LENGTH (MM)

Figure 4. Histograms of length in Laccophilus maculosus. Males are shown
crosshatched; females, stippled.

since an intergrade should have it situated in a position intermediate
between those of decipiens and shermani. Only a few females from
the intergrade zone have the flange; and, usually, the sample size is
too small to collect one with a flange.

Both intergrade populations show chaotic mixing; thus, it seems
that, in both cases, secondary hybridization has occurred. Within the
zone, any one specimen may 1) show no sign of intermediacy, 2)
show only a trace of mixture in perhaps but one character, or 3) be

MEM. AMER. ENT. SOC., 26

62 THE GENUS LACCOPHILUS

intermediate in all characters (rarely). Elytral patterns or aedeagi
provide the best evidence for intergradation, but the coxal files are
also useful on occasion. A population from El Paso County, Colo-
rado, which appears to be well within the range of decipiens, con-
tained two specimens that had only 36 lines in the file; five others had
counts in the middle forties, which is more typical of decipiens.
Apparently, there is some gene flow from shermani influencing the
population.

The narrowness of the zones is difficult to explain. Possible ex-
planations are that there is incomplete gene flow between the races
or that the ecological requirements of the races are sufficiently strin-
gent to limit the intergrades to an intermediate environment. The
latter appears true in the case of decipiens and maculosus, since they
merge in the northern plains — beyond their usual habitats of ponds
in coniferous or deciduous forests. If one assumes that the range of
the two races has been expanding and that permanent populations are
relatively scarce in the area of intergradation, then it might be ex-
pected that there would be considerable variation in the degree of
hybridization in different populations.

This argument is less pertinent to decipiens and shermani, how-
ever. There is little or no apparent ecological difference on either
side of the intergrade zone, but the zone appears extremely narrow —
perhaps not over 100 miles wide. More analysis may show a wider
zone than indicated in the present paper, but as yet there is no evi-
dence of swamping. It is entirely possible that an equally correct
argument could be presented for maintaining shermani as a separate
species.

MATERIAL EXAMINED
L. m. maculosus X L. m. decipiens

CANADA. — MANITOBA. Aweme, 1, vi.4.03, N. Criddle (USNM); 1 4,
1 9,.1x.1.28, E. C. (USNM). MHusavick, 2 6, 2 9, vii.8.—, J. B. Wallis
(AMNH). Man., 1 2 (MCZ). Stony Mtn., 1 2, v.28.—, J. B. Wallis (AMNH).
Treesbank, 9 6, 5 9, vii.25.—, J. B. Wallis (AMNH). Winnipeg, 1 6, viii.27.27,
P. J. Darlington, Jr. (MCZ); 1 6, iv.23.25, L. H. Roberts (USNM).

UNITED STATES OF AMERICA. — COLORADO. Boulder County. Boul-
der, 1 2, ix.25.17, Hubbard; 1 2, vili—.16, S. D. Pokling (AMNH). Larimer
County. Ft. Collins, 1, x.15.47 (USNM); 1, x.8.51, Bryant; v.20.32; County,
1 ¢ (AMNH). NEBRASKA. Thomas County. Dismal River on Hwy. 83,
19, vii.15.67, H. B. Leech (CAS). SOUTH DAKOTA. Custer County. Custer

JAMES R. ZIMMERMAN 63

St. Pk., 1 2, vi.20.55, W. A. Drew (MCHS). Pennington County. Badlands
N. Mon., 1 9, viii.7.52, B. Malkin & V. Thatcher (FM). Wasta, 1 ¢, viii.7.52,
B. Malkin (FM).

L. m. decipiens X L. m. shermani

UNITED STATES OF AMERICA. — ARIZONA. Navajo County. Little
Colorado Riv., 8 ¢6, 8 2, vii.23.63, PSB (CAS). CALIFORNIA. Riverside
County. County, Sonorian (sic!) Region, Colorado R., 10 6, 4 2, F. E.
Winters; 8 4 with shermani aedeagus, 1 4 with decipiens aedeagus and 1 4 an
intergrade; all females, with epipleural expansion in shermani position. NEW
MEXICO. Colfax County. Cimarron, 10 m. N., 2 6, 8 9, vii.15.65, A. H.
Smith & T. O. Boswell (NMSU). Rio Arriba County. Canjilon, 3 m. W.,
1 6, 5 9, ix.25.65, A. H. Smith & T. O. Boswell (NMSU). El Rito, 18 ¢,
13 2, vi.23.65, JRZ, A. H. Smith, R. L. Smith; 9 ¢, 19 9, ix.25.65, A. H.
Smith, T. O. Boswell (NMSU). Brazos, 1 2, viii.11.65, H. B. Leech (CAS).
Ojo Caliente, 44 6, 27 2, vi.23.65, R. L. Smith, A. H. Smith, JRZ (NMSU).
Parkview (11 m. S., Chama), 8 é, 9 2, vi.23.65, JRZ, A. H. Smith, R. L.
Smith (NMSU). San Miguel County. Las Vegas, 21 m. S., 1 2, vii.16.65,
A. H. Smith & T. O. Boswell (NMSU). Mosquero, 19 m. W., 3 6, 12 92,
x.23.65, A. H. Smith (NMSU). Trujillo, 19 m. E., Trementina Ck., 4 6, 6 2,
A. H. Smith & T. O. Boswell (NMSU).

LACCOPHILUS FASCIATUS

This is a wide-ranging polytypic species with three readily dis-
tinguished subspecies. Each race was previously considered a sep-
arate species. Laccophilus fasciatus (= apicalis Sharp) was described
by Aubé in 1838 from material from the United States and Mexico.
The only type still available is from Mexico. L. rufus was described
by Melsheimer in 1846 from Pennsylvania and was long considered
to be a synonym of L. fasciatus. L. terminalis is from Texas (Sharp
1882a). The nominate subspecies is a tropical one that extends south
in South America and north to the Mexican Plateau. It intergrades
with terminalis in the northeastern and northwestern coastal lowlands
of Mexico. The latter occurs throughout the Mexican Plateau, the
southwestern United States, and the Great Plains to South Dakota.
It intergrades with rufus from Kansas (or Nebraska?) south to east-
central Texas. L. f. rufus occurs throughout the eastern United States
north of the Florida Peninsula.

L. f. terminalis noticeably differs from rufus only in elytral pat-
tern and aedeagus. The former may be slightly larger, but neither
shows any differences in size between the sexes. L. f. fasciatus dif-

MEM. AMER. ENT. SOC., 26

64 THE GENUS LACCOPHILUS

fers from terminalis, not only in elytral pattern and aedeagus, but
also in quantitative characters. It is smaller, relatively wider, and
has a difference average size for males over females.

Intergradation between rufus and terminalis has resulted from
secondary contact at about the boundary of eastern acid pedalfer soils
and western alkaline pedocals. One race is the geographical and
ecological replacement of the other, and intergrades persist in an in-
termediate environment. The pattern of intergradation is complex
between fasciatus and terminalis. Intergrade populations commonly
occur in the coastal lowlands and lower mountains from Nayarit to
southern Arizona and, more rarely, (perhaps due to insufficient col-
lecting) in the northeastern Mexican coastal lowlands. In Jalisco,
however, fasciatus and terminalis are sympatric. This situation is
confusing, since there appears to be no barrier between them and
intergrading populations in Nayarit.

DESCRIPTION. — Medium-sized (length 4.3 to 5.2 mm; width 2.3 to 2.8
mm), brown, irrorated species with more or less complete black fascia or blotch
across the posterior half of elytra; metacoxal file present; prosternal process
short; ovipositor sawlike. COLOR. Head: yellow above and beneath, some-
times with reddish-brown tint, especially between the eyes at the base of the
pronotum; appendages pale yellow except mandibles which darken to brewnish-
red toward tip. Pronotum: pale brownish-yellow with reddish tint. Elytra:
brown to dark brown irrorations on a pale yellow or brownish-yellow back-
ground, intensifying and coalescing in the posterior half to form a transverse
fascia or blotch that varies from merely suggested in terminalis to solid black
from one side to the other in fasciatus (figs. 282, 288); anterior half with
finger-like extensions irroration of variable intensity and completeness; apices
pale or darkened; epipleura pale yellow in broad anterior half, but dark brown
in constricted posterior half. Tergite VIII: basal half dark brown; distal half
of the same color in the nominate race, but pale yellow or reddish-yellow in the
other two races. Venter: varying from yellow to reddish-brown; usually dark-
est on lateral portions of metacoxal plates and posterior margins of abdominal
sclerites; front and middle legs pale yellow; hind legs yellow basally and be-
coming more reddish-brown distally; oblique striations variable from yellow to
reddish-brown. Genitalia: variable from yellow and light brownish-red to dark
reddish-brown; oval plate and left paramere usually paler than other parts.
ANATOMY. Méicroreticulation: weakly double on head, pronotum, and elytra;
secondary mesh discernible, but individual cellules still apparent. Head: supra-
clypeal seam closely parallel to margin. Pronotum: WH/PW, 0.67 to 0.69;
LP/PW, 0.41 to 0.42. Elytra: epipleural flange frequently present in females
of the nominate race, but absent in the other females; truncation of the elytral
apices slight. Venter: coxal file prominent in males, composed of about 20 to
30 lines; prosternal process with well-defined crest; lobes of postcoxal processes

65

JAMES R. ZIMMERMAN

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MEM.

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6100°0 L9S'0 6S0';0 IIc c60'0 + S9'C (OO1) TITO TLE Isto OLY Tel “OD YOIMSpPIG
cC00'0 999°0 c90'0 CIS LLOO S9°C (€r) 6600 lt IcloO SLY 19 sesuvy “OD AsareH
ou) Td/Md yIPIM qipIM (N) yisua| yysu9| N Ay e907]
jejouo1d yeayAjo

syounudsa) “fT «x snf{na *f °7

(panuljuor) Z 2QeL

AMER. ENT. SOC., 26

MEM.

70 THE GENUS LACCOPHILUS

rounded and laterally projecting well posterior to midline; last visible ventral
abdominal segment rounded and similar in males and females; that of males
slightly produced and with an asymmetrical curving ridge in middle; last seg-
ment of females weakly arched with posterior margin not noticeably produced.
Legs: male pro- and mesotarsi enlarged in a dorsoventral plane; palettes easily
distinguished at 20 power magnification; male fifth tarsal segments on both pair
of legs twice as long as corresponding fourth; profemoral setae (5 to 7) shorter
and finer than mesofemoral ones (5 to 7). Genitalia: oval plate large and pro-
duced to acuminate tip with ventral crest curving to the right as it extends for-
ward; an indefinite number of raised lines lying on either side of crest; aedeagus
curved and narrowing toward apex; right paramere produced at apex; ovipositor
with about 11 teeth on each ridge.

Laccophilus fasciatus fasciatus Aubé, new synonymy and new status
(Figs. 39-44, 47, 50, 51, 288)

Laccophilus fasciatus Aubé, 1938, p. 423. Type: Institut Royal des Sciences
Naturelles de Belgique, number 2842, female: type locality here restricted
to Cuitlahuac, Vera Cruz, Mexico.
Laccophilus apicalis Sharp, 1873, p. 53; Sharp, 1882a, p. 291; Sharp, 1882b,
p. 10; Darlington, 1936, p. 153; Leech, 1948a, p. 65.

DIAGNOSIS. — The dark blotch across the posterior part of the elytra will
separate this race from all other irrorated species in Mexico and Central Amer-
ica except salvini. It is black beneath, however, while fasciatus is reddish-
brown to brown. L. salvini also lacks subbasal finger-like projections on the
elytral pattern.

NOMENCLATURAL NOTES. — This race has long been rec-
ognized as L. apicalis Sharp, and fasciatus has been ascribed to the
populations in the eastern United States; but, unfortunately, Aubé
described fasciatus from mixed material from the United States and
Mexico. The only known specimen used in the original description is
a female from Mexico; thus, the name fasciatus must be used for the
southern subspecies. The species name remains the same, since both
are members of one polytypic species.

VARIATION. — Individual variation in color and pattern seems
as great as any geographic variation except for intergrade influence
from terminalis. There are some important sexual and geographic
variations in quantitative characters, however. L. f. fasciatus males
are about 0.1 mm longer and 0.05 mm wider than females (Table 2).
The PW/EL ratio, which is the same for both sexes, shows an in-
crease in Honduras and Costa Rica. From Nicaragua north, the
PW/EL value is about 0.583 with no value exceeding 0.587; but
south of Nicaragua, the averages are 0.587 to 0.596. This means

JAMES R. ZIMMERMAN 71

|
LACCOPHILU F. TERMINALIS

LACCOPHILUS F. FASCIATUS

Figure 5. Distribution of Laccophilus fasciatus.

that the southern Central American populations are relatively wider
than the ones to the north. There does not appear to be any significant
difference in length, however, between the two areas.

But specimens from the Yucatan Peninsula are larger. A sample
(N = 46 for each sex) from Chenchoyi, Campeche, was larger in all
measurements than any other locality except Autlan, which might be
reflecting influence from terminalis. The closest samples to Chen-
choyi (Catemaco, Vera Cruz and Chiapa de Corzo, Chiapas) are
smaller for all measurements, although the WP/EL ratios are the same.

The heterogenity of L. f. fasciatus is also reflected in a plot of the
frequency distribution for length (Fig. 6). The curve for females
has two peaks — one at about 4.75 mm and the other at 4.60 mm.

MEM. AMER. ENT. SOC., 26

V2 THE GENUS LACCOPHILUS

terminalis

NUMBERS

4.2 4.4 4.6 4.8 5.2

fasciatus

5.2

TOTAL LENGTH (MM)

Figure 6. Histograms of length in Laccophilus fasciatus. Males are shown
crosshatched; females, stippled.

The number of specimens is too large to assume sampling error in
this case. There is only one peak for males (also at about 4.75 mm),
but there is a bulge on the left slope which suggests another peak
might be hidden in the curve. In contrast, the curves for terminalis

JAMES R. ZIMMERMAN 73

and rufus have single definite peaks and even slopes. The larger pop-
ulation from the Yucatan Peninsula might account for these
irregularities.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— The northern boundary of L. f. fasciatus forms a U-curve lying on
the separation of the Nearctic and Neotropical Realms. It inter-
grades with terminalis in coastal Tamaulipas in northeastern Mexico
and in several states in northwestern Mexico. Both races occur in
Jalisco, but evidence of intergradation is slight. L. f. fasciatus ap-
pears to be continuously distributed in the lowlands all the way into
South America and is generally absent from the volcanic highlands of
Puebla, Morelos, Mexico, and Guerrero.

It is usually found below 3000 feet. It is most common in road-
side ditches, pools of streams, sloughs, and stock ponds. The race is
most abundant in shallow water in marginal vegetation and debris.

MATERIAL EXAMINED

COSTA RICA. — Bagaces, Guanacaste, 6 ¢, 5 @, vii.12.57; Guanacaste, 1 é,
2 2, vii.17.57, D. R. Lauck (USNM). La Cruz, 16 m. S., 100+, vii.25.65;
Liberia, 8 m. SE., 10 6, 12 2, vii.24.65; Pacuare, 2 2, vii.2.67; Punta Arenas,
2 6, 4 Q, vii.22.65; Rivas, 22 m. S., 2 2, vii.26.65, P. J. Spangler (USNM).
San Jose, 2 6, 2 9, vii.16.57, D. R. Lauck (USNM). Taboga, Agr. Exp. Sta.,
34 6, 40 9, vi.27.67; 14 6, 7 2, vi.28.67; Turrialba, 50+, vii.15.65, P. J.
Spangler (USNM).

EL SALVADOR. — Acajutla, 19 m. S., 3 6, 2 9, vii.8.65; Acajutla, 7 ¢, 7 2,
vill.4.64; Cd. Arce, 7 m. SE., 1 4, viii.3.65; La Union, 15 m. SE., 14 4, 10 @,
vil.31.65, P. J. Spangler (USNM). La Union, Santa Rosa, 1 2, vii.23.57; 2 ¢,
vii.24.57, D. R. Lauck (USNM).

GUATEMALA. — Aldea Jesus Maria, 49 ¢, 54 92, viii.15.65; El Progreso,
5m. S., 2 2, vili.11.65; Escuintla, 17 m. E., 2 ¢, vii.8.65; Guatemala City, 20
m. S., 1 6, 1 2, vil.7.65, P. J. Spangler, Jr. (USNM). Mazatenengo, 1 2,
vili.1.57, D. R. Lauck (USNM). Morales, 1 m. N., 1 6, viii.16-18.65; Pijije,
26,2 2, vil.8.65; 15 m. W., 4 6, 10 Q, viii.5.65, P. J. Spangler, Jr. (USNM).
Suchit, Rio Sis, 22 km. S. La Maquina, 1 ¢@, vi.11.65, O. S. Flint (USNM).

HONDURAS. — Choluteca, 10 m. W., 25 +, vii.29.65; Jicaro Galan Jct., 2 2,
vii.29.65, P. J. Spangler, Jr. (USNM). Morazan, 5 6, 4 2, vil.22.57, D. R.
Lauck (USNM). Pespire, 2 ¢, 2 9, vii.29.65; Sabana Grande, 10 m. N., 4 6,
10 Q, vii.29.65, P. J. Spangler, Jr. (USNM).

MEXICO. — CAMPECHE. Champoton, 11 m. SW., 7 6, 3 2, xi.27.63; Chen-
cholli, 100 6, 122 2, xi.28.63; Hopelchen, 18 m. NE., 6 6, 11 2, x1.28.63, JRZ
(NMSU). CHIAPAS. Chiapa de Corzo, 4 m. E., 27 6, 26 9, 1x.1.63; Cin-

MEM. AMER. ENT. SOC., 26

74 THE GENUS LACCOPHILUS

talapa, 5 m. W., 2 6, 4 9, ix.1.63; Comitan, 8 6, 13 92, viii.30.63; 13 m. W.,
1 6, 2 9, vili.30.63; Cuahtemoc, 5 6, 6 9, viii.30.63; 4 m. W., 16 6, 18 2,
vili.30.63, JRZ (NMSU). Las Cruces, 5 m. S., 12 6, 17 9, viii.23.65; P. J.
Spangler, Jr. (USNM); Near Pichucalco, 3 6,2 2, xi.26.63; Ocozucuatla, 6 m.
W., 4.4, 3 2, ix.1.63; Ixtapa, 1 ¢, viii.31.63; Tuxtla Guttierrez, 2 6, 2 2,
ix.1.63; 10 m. W., 5 6, 3 Q, ix.1.63; JRZ (NMSU). COAHUILA. Torreon,
1 6, M. Embury (CAS) (Intergrade?). COLIMA. Colima, 7 m. NE., 6 6,
2 2, xii.3.48, H. B. Leech (CAS). Manzanillo, 5 m. S., 4 6, 3 2, vii.29.62;
Trapechi, 9 6, 4 9, vii.30.62, JRZ (NMSU). JALISCO. Atenquique, 1 2,
xii.5.48, H. B. Leech (CAS). Autlan, 9 m. SW., 34 6,55 2, x.24.66; Autlan,
17 6, 18 9, x.26.66, A. H. Smith & JRZ (NMSU). Barra de Navidad, 1 3,
6 @, ili.18.61, C. O. Morse (CAS); 2 6, 2 2, vi.6.63, K. L. McWilliams; 5 é,
8 2, x.25.66; La Heurta, 6m. S., 1 6, 1 9, x.24.66;5 m.N., 7 4, 7 9, x.25.65,
A. H. Smith & JRZ (NMSU). Mazamitla, 1 m. N., 1 4, 2 9, ii.8.53, I. J.
Cantrell (UMMZ). Between Union de Tula and Autlan at Rio de San Pedro,
22 6, 34 Q, iii.28.64, JRZ (NMSU). Near Atenquique (29 m. NE. of Colima
by road), 7 ¢, 23 9, xii.3.48, H. B. Leech; this locality is given as Colima, but
must be in Jalisco (CAS). Tehetlican, Rd. 10, 2 4, vii.28.62; 10 m. S., 2 6,
3 2, JRZ (NMSU). MICHOACAN. Urupan, 10 m. S., 1 4, iii.26.64, IRZ
(NMSU). OAXACA. Juchitan, 3 m. E., 27 6, 17 @, ix.7.64; 20 km. E., 2 4,
4 2, ix.7.64; 15 m. of Juchitan on Rd. 185, 2 ¢, 4 @, ix.2.63, JRZ (NMSU).
La Ventosa, 50 m. N., 1 ¢@, xii.14.55, J. C. Schaffner (UMMZ). Matias
Romero, 5 m. S., 1 @, ix.7.64, JRZ (NMSU). Oaxaca, 4 6, 4 2, vii.20.37,
H. B. Leech (CAS). Salina Cruz, 5 ¢, 5 9, 1x.6.64; Tapanatepec, 17 6,
18 9, ix.1.63, JRZ (NMSU). Tehuantepec, 17 ¢, 25 9, 1x.2.63; 4 4, ix.6.64;
Texquisixtian (Rd. 190), 5 6, 8 2, JRZ (NMSU). Temascal, 1 ¢, 1 2,
v.25.64 (CAS). PUEBLA. Near Maria Andrea, 3 6, 1 2, ix.10.64, JRZ
(NMSU). TABASCO. Teapa, 5 m.N., 1 2, ix.2.63, JRZ (NMSU). VERA-
CRUZ. Acayucan, 1 2°, vili.6.57, D. R. Lauck (USNM); 20 m. S., 4 6, 2 2,
ix.7.64; 10 m. E., 2 9, ix.7.64; Catemaco, 5 6, 5 2, viii.26.62; Cuitlahuac,
17 6, 13 9, viii.25.62, JRZ (NMSU); 300+, viii.10-12.64, P. J. Spangler, Jr.
(USNM). Huatusco, 25 km. E., 3 6, ix.9.64, JRZ (NMSU). Jalapa, 10 m.
E., 2 2, vili.27.62; J. D. Covarrubia, 1 m. N., 4 6, viii.26.62; Martinez de la
Torre, 19 6, 9 9, ix.9.64; Near La Tinaja, 8 ¢, 28 9, vili.25.62; Near Garro,
2 6, ix.8.64; Papantla, 18 km. E., 3 é6, 4 2, ix.9.64; Paso del Toro, 3 2,
vili.25.62; 15 km. W., 1 é&, ix.8.64; Paso de Ovejas, 3 ¢, 8 2, vili.27.62, JRZ
(NMSU). Poza Rica, 9 m. W., 5 4, 3 2, viii.27.65, P. J. Spangler, Jr.
(USNM.) Puenta Jula, 3 ¢, 13 2, xii.18.48, H. B. Leech (CAS). Santiago
Tuxtla, 10 km. S., 2 6, ix.8.64; Tecolutla, 15 km. S., 1 2, ix.10.64, JRZ
(NMSU). Veracruz, 1 ¢, H6ge (AMNH); 2 9, vii—.59, N. Krauss (CAS).
YUCATAN. Uxmal, 2 6, 1 9, xi.25.63, JRZ (NMSU).

NICARAGUA. — Belen Rivas, 4 6, 5 92, vii.18.57, D. R. Lauck (USNM).
Esteli, 3 ¢, 2 9, vii.31.67, O. S. Flint (USNM); 9 m. N., 50 6, 50 9, vii.10.65;
La Trinidad, 100+, vii.27.65, P. J. Spangler, Jr. (USNM). Madriz, 6 6, 4 2,
vii.8.57, D. R. Lauck (USNM). Rivas, 10 m. N., 100+, vii.11.65; San
Benito, 13 m. N., 34 6, 38 9, vii.11.65, P. J. Spangler, Jr. (USNM). San

JAMES R. ZIMMERMAN is)

Benito, Managua, 2 ¢, 1 2, vii.10.57, D. R. Lauck (USNM). Somoto, 100+,
vii.28.65, P. J. Spangler, Jr. (USNM).

PANAMA. — Algarrobos, 9 m. S.,9 ¢, 10 2, vil.5.67; Anton, 5.3 m. E., 15 6,
17 @, vii.6.67, P. J. Spangler, Jr. (USNM). Playa San Carlos, 3 ¢, vii.15.67,
O. S. Flint (USNM).

Laccophilus fasciatus rufus Melsheimer, restored name and new status
(Figs. 39-45, 49, 57, 284, 285)
Laccophilus rufus Melsheimer, 1844, p. 28. Type: Harvard University, Mu-
seum of Comparative Zoology, number 31234, male, Pennsylvania.

Laccophilus fasciatus, Crotch, 1873, p. 400; Sharp, 1882a, p. 290; Blatchley,
1910, p. 210; Leng, 1920, p. 77; Zimmermann, 1920, p. 18; Zimmerman
and Severin, 1957, pp. 29-32; Zimmerman, 1960, pp. 142-3.

DIAGNOSIS. — L. fasciatus can easily be separated from all other irro-
rated species in the eastern United States by the fascia or blotch on the posterior
half of the elytra.

NOMENCLATURAL NOTES. — Say (1823) recognized rufus,
but referred to it as a variety of maculosus without giving it a name.
This race has long been known as fasciatus; but, as discussed above,
the remaining type is the Mexican subspecies and not an individual
from the eastern United States. Melsheimer’s rufus is restored as
valid, since there is no doubt as to the identity of his type in the
Museum of Comparative Zoology.

VARIATION. — The principal variation in elytral pattern is
shown in Figures 285 and 286. Small specimens with weakened fas-
cias resemble the sympatric proximus. This is individual variation,
however; and, apparently, has no geographical significance.

Males and females are very nearly the same size, both averaging
about 4.74 mm; and measurements have been combined for each
locality.

Apparently, there is considerable regional variation in size of
rufus, while pattern and the PW/EL ratio remain fairly uniform
throughout the range.

The average values for all four measurements are much the same
for three large samples from Indiana, Illinois, and Kansas (Table 2),
but three other samples from the eastern United States give a different
picture. Sixty-nine specimens from several southeastern states had a
mean length of only 4.68 mm, as compared to 4.73 mm for the three
midwestern samples and 4.90 mm and 4.81 mm for New York and

MEM. AMER. ENT. SOC., 26

76 THE GENUS LACCOPHILUS

Pennsylvania. L. f. rufus appears to be smaller in the southeast than
in any other part of the range. The standard deviations were all
larger than other comparable samples, which indicates some currently
unrecognized difference within southeastern populations.

A cline for all four measurements shows up if the three samples
from New York to Indiana are compared. The differences are all
significant (95 per cent level and above). This is one of the few
clines that have been discovered in this study even though the popu-
lations cover large areas.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— L. f. rufus ranges from northern Florida and east Texas to Ver-
mont, southern Michigan, and southeastern South Dakota and from
the Atlantic Coast to a line approximating the ninety-seventh meridian
in South Dakota, Nebraska, Kansas, Oklahoma, and Texas. It nor-
mally does not occur in the peninsula of Florida, or the lowlands
bordering the Gulf of Mexico; but it has been taken in the vicinity of
Mobile Bay. It is probably the most abundant Laccophilus south of
the glaciated region in the eastern United States. It intergrades with
terminalis from northcentral Kansas to east Texas.

Young (1954) found rufus most commonly in muddy or silty bot-
tomed temporary pools formed in roadside ditches or intermittent
streams. These pools were in well-drained soils. It was a pioneer
species of recently formed aquatic habitats in upland Florida. Zim-
merman (1960) found it in open, unshaded ponds and roadside
ditches in clay soils in the Midwest.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ALABAMA. Mobile County. Mobile,
8, J. D. Sherman, Jr. (USNM). Montgomery County. Barachias, 2, vi.10.24,
E. G. Holt (USNM). Russell County. Seale, 1 9, vi.18.54, R. L. Fisher,
D. L. Haynes (MCHS). ARKANSAS. Grant County. Poyem, 10 6, x.25.48,
O. Bryant (CAS). Hempstead County. Hope, 1, ix.7.31, Knoble (BERK);
4 2, iv.3.22, L. Knoble (CAS); 1 4, vi.l1.54, J. W. Green (CAS). CON-
NECTICUT. Fairfield County. County, 1 2, viim—.—, C. H. Roberts (AMNH).
DISTRICT OF COLUMBIA. Piney Br., 1, iv.25.05 (USNM); 1, viii.26.06,
D. H. Clemons (USNM). Rock Creek, 6, vili.31.06, D. H. Clemons (USNM).
Washington, 1, vii.17.—; 1, viii.28.—, Hubbard & Schwarz (USNM); 3, ix.7-
23.06, W. L. McAtee (USNM); 1, vi.6.04, F. Knab (USNM); 1, vii.31.—,
H. S. Barker (USNM); 3, iv.28.18, L. L. Buchanan (USNM). _ District 4,
—.—.— (USNM). FLORIDA. Duval County. Jacksonville, 1 6, Van Dyke
(CAS). GEORGIA. Chatham County. Savannah, 1, v.17.27, E. W. Mank

JAMES R. ZIMMERMAN il

(CNL). Cobb County. Marietta, 2, iii.5.11 (CNL); 1, iii.5.11 (USNM).
Decatur County. Bainbridge, 1 6, 1 2, vii.15-27.09, J. C. Bradley (CAS); 6,
vii. 15-27.09; 1, iii.7.10, J. C. Bradley (CNL); County, 2, x.3.38, F. N. Young
(CNL). Jefferson County. Wrens, 2, iii.8.11 (CNL); 1, iii.8.11 (USNM).
Rabun County. Clayton, 1 4, vii—.10, W. T. Davis (CAS); County, 4 é,
3 2, vii .—, C. V. Leng (CAS). ILLINOIS. Alexander County. Olive
Branch, 8 6, 11 2, x.8.09, Gerhard (FM). Bond County. Mulberry Grove,
6 6, 3 2, x.20.56, JRZ (NMSU). Champaign County. Urbana, 1, vii.19.07
(USNM); 1 4, 1ii—.— (CNG); Champaign, 2 6,5 2, xi.5.27, A. T. McClay
(DAV). Cook County. (Morton Gr.) Edge Brook, 2, vii.15.11, J. D. Sher-
man, Jr. (USNM). Glencoe, 2, 1x.5.04, J. D. Sherman, Jr. (USNM). Palos
Pk., 1 2, x.14.34, W. J. Gerhard (FM). Franklin County. Zeigler, 1, iti.13.29,
J. Karlovich (USNM). Kankakee County. Momence, 1, vii.—.—, Wickham
(USNM). Jackson County. Murphysboro, 3 6, 1 9, viii.9.32, W. J. Gerhard
(FM). Pike County. Pittsfield, 3 3, viii.17.46, 1 6, 2 2, viii.7.47; 1 ¢,
Vili.22.47; 1 2, vii.28.48, B. Cadwell (DAV). Randolph County. Bremen, 1,
vii.19.56, P. J. Spangler, Jr. (USNM). Vermilion County. Oakwood, 1 é,
v.5.48, H. T. McClay (DAV). INDIANA. Brown County. Brown County St.
Pk., 8 6, 7 9, iv.21.56, JRZ (NMSU). Clinton County. Kirklin, 3 m. W.,
il @5 I Ss mele il @, il 2, De POSES © 6, 6 25 so IO SSS 7 G5 Bp xa) 7s
JRZ (NMSU). Dubois County. Purdue Exp. Farm, Near Huntingburg, 5 ¢,
5 2, x.29.55, J. E. Hanegan & JRZ (NMSU). Hamilton County. Westfield,
7m. N., 3 2, v.4.56, JRZ (NMSU). Huntington County. Huntington, 2 m.
N., 7 6, 9 2, x.26.56, JRZ (NMSU). Knox County. Hazelton, 10 m. W.,
2 6,2 2, 1x.9.56, JRZ (NMSU). Lake County. Clarke, 2 2, viil.7.07, (FM).
Madison County. Alexandria, 5 m. E., 4 6, 3 2, vii.10.57, JRZ (NMSU).
Anderson, 2 92, vi.6.57;2 6, vi.21.57,JRZ (NMSU). Monroe County. Bloom-
ington, 12 6, 12 9, ix.15.55; 6 6, 10 @, ix.20.55; 19 6, 4 9, 1.13.56; 22 6,
29 2, iv.26.56; 1 6, 6 2, v.11.56; 13 6, 15 9, vii.1.56, JRZ (NMSU); 4 m.
N., Slough, 12 ¢, 8 2, vi.18.57, JRZ (NMSU); 4 6, 7 2, xii.31.49, F. N.
Young (AMNH); 4, xii.31.49, F. N. Young (CNL); 1.9.52, F. N. Young
(CNL); 1 6,1 9, xii.31.49, F. N. Young (FM); 1 ¢, 1 2, same data (ANSP).
Morgan County. Waverly, 5 2, vi.17.57; Morgan-Monroe St. For., 1 ¢,
vii.15.57, JRZ (NMSU). Noble County. Wolflake, 5 m. S., 1 é, x.26.56,
JRZ (NMSU). Owen County. McCormick’s Ck. St. Pk., 3 6, 2 2, vii.12.55;
Richland Ck., Near Whitehall, 2 ¢, 2 2, viil.26.55, JRZ (NMSU). Posey
County. Mt. Vernon, 5 m. E., 4 3, 7 2, ix.9.56, JRZ (NMSU). Wabash
County. Wabash, 3 m. N., 3 6, x.26.56, JRZ (NMSU). IOWA. Decatur
County. Leon, 1, vii.3.32, F. Andre (TAM). Lee County. Fort Madison,
12 (AMNH). Polk County. Herrold, 1, vii.24.19, E. D. Quirsfeld (CNL).
Story County. Ames, 1, v.11.32, Floyd Andre (USNM). KANSAS. Atchin-
son County. Atchinson, 1 é, ix.22.56, J. W. McReynolds (CAS). Cherokee
County. Galena, 19 (CNL); 19 (USNM); 33, viii.26.04 (USNM); 6 4, 11 @,
Ebb. Crumb (AMNH). Douglas County. Baldwin (City?), 3 6, 4 2, J.C.
Bridwell (ORES). Lawrence, 1, x.9.49, J. G. Rozen (BERK): 23 4, 25 9°,

MEM. AMER. ENT. SOC., 26

78 THE GENUS LACCOPHILUS

ix.26.—; x.2.51, P. J. Spangler, Jr. (USNM). Lone Star L., 15 4, 13 2,
vi.6-10.52, P. J. Spangler (USNM). County, 2, x.9.48, J. G. Rozen (BERK);
20, 11.20.52; 3, iti.19.52; 1, iii.11.52, P. J. Spangler, Jr. (USNM); 2, F. H.
Snow (CNL). Leavenworth County. Lawrence, 7 m. NE, 2, iii.22.52, P. J.
Spangler, Jr. (USNM). Ottawa County. County, 1 ¢, iv.31.33, C. W. Sa-
brosky (MCHS). Pottawatomie County. Onaga, 2 (USNM); Onega (Onaga?),
1 6, 2 2, F. Crevecoeur (AMNH). Riley County. 1, iii.13.—; 1, viii.1.—;
11, x.21.—, Popenoe (USNM). Shawnee County. Topeka, 4 (USNM); 3,
Hubbard & Schwarz (USNM); 2 2, Popenoe (MCZ). LOUISIANA. Natchi-
toches County. Coverdale (AMNH); 2 (USNM). Vowell’s Mill, 7 6, 9 2
(CAS). Vernon County. Leesville, 1 2, viii.18.47, H. S. Fitch (DAV).
MARYLAND. Baltimore County. 1 6, vii.13.—, F. E. Blaisdell (CAS).
Harford County. Edgewood, 2, ix.17.18, H. Dietrich (CNL). Montgomery
County. Kensington, 36, vi.2-17.61, J. W. Fitzgerald (USNM). Piney Point,
3, vili.26.46, R. I. Sailer (USNM). Plummers Island, 3, v.6.05; 1, iii.24.07;
3, 1v.27.13, W. L. McAtee (USNM). MICHIGAN. Van Buren County.
South Haven, 1, viii.11.04 (USNM). Wayne County. Detroit, 2, vi.28.—,
2, Hubbard & Schwarz (USNM). MISSISSIPPI. George County. Lucedale,
5, 1.17.32; 4, iii.27.32, H. Dietrich (CNL). Perry County. Beaumont, 1,
iv.19.32, H. Dietrich (CNL). Wilkinson County. Woodville, 1 4, vii.26.21
(CAS). MISSOURI. Boone County. Ashland, 1, iii.29.56, P. J. Spangler, Jr.
(USNM). Columbia, 10, iii.21-29.36; 3, v.2.36, W. M. Gordon (CNL); 3,
vi.26.—, Wickham (USNM). Columbia, 5 m. S., 1, viii.20.53, M. C. Grabeau
(USNM). Deer Park, 6 m. E., 1, viii.28.53, M. C. Grabeau (USNM). Cooper
County. Booneville, 1 m. E., 6, ix.25.54, P. J. Spangler, Jr. (USNM). Gi-
rardeau County. Cape Girardeau, 1, iv.1.56, D. Stout (USNM). Greene
County. Springfield, 1, xii.11.49, E. T. Oxtman (BERK). Mississippi County.
Charleston, 1, iv.23.56, P. J. Spangler, Jr. (USNM). St. Louis County. Howard
Bend, 7, vi.28.37; 7, vii.17.37; 2, vii.22.40; 1, ix.18.40, W. M. Gordon (CNL).
Manchester, 1, 1x.15.57, P. J. Spangler, Jr. (USNM). St. Louis, 1, M. Schuster
(USNM). Webster Grove, 1, vi.27.33, R. C. Lange, (USNM). Vernon
County. Near Nevada, 1 ¢, 3 9, v.14.58, J. W. McReynolds (CAS). Nevada,
16, 1 9, vi.l1.58, J. W. McReynolds (CAS). County, 1 2, v.30.58, J. W.
McReynolds (CAS). NEBRASKA. Lancaster County. Lincoln, 2 6, 3 9,
x.15.60, W. F. Rapp, Jr. (NMSU). NEW JERSEY. Bergen County. Fort
Lee, 7, vii.30.04, J. D. Sherman, Jr. (USNM). Palisades (Pk.?), 1, iv.19.03,
T. D. O'Connor (CNL). County, 4, v.11.02, Shoemaker (USNM). Camden
County. Delair, 3 46, 4 2, vii.16.—, E. L. Dickerson (AMNH). Middlesex
County. Avenel, 1 2, v.22.31, Siepmann (FM). Spotswood, 1 2, C. H.
Roberts (AMNH). Morris County. Boonton, 1, vii.8.01, G. M. Greene
(USNM). Ocean County. Ft. Dix, 1 &, vi.10.43, C. D. Michener (AMNH).
Lakehurst, 1, vi.26.54, D. M. Anderson (CNL); 1, ix.1.01, J. D. Sherman, Jr.
(USNM); 4 6, 3 9, vii.17.23, L. B. Woodruff (AMNH); 1 2, v.24.—; 1 ¢,
vii.31.—, C. H. Roberts (AMNH). NEW YORK. New York Vicinity. Staten
Is., 1 6, Leng, Van Dyke (CAS); 1; 1, vii.—.91; 17, 1x.15.00, J. D. Sherman,
Jr. (USNM): 1 9, ix.27.38; 1 2, x.1.38; 1 9, i1x.4.39; 1 2, v.14.39; 1 6, 1 9,

JAMES R. ZIMMERMAN 719

iv.23.39 (FM); 2 9, iv.19.—, C. H. Roberts (AMNH); 1 2, ix.27.38 (DAV).
N.Y.C., 1 ¢, 1 29, C. H. Roberts; 1 ¢, x.24.01 (AMNH). Westchester County.
Peekskill, 1, J. D. Sherman, Jr. (USNM). NORTH CAROLINA. Buncombe
County. Ashville, L. Ashnoca, 1, iv.29.44, W. E. Hoffman (USNM); 1, v.—.44,
W. E. Hoffman (USNM). Ashville, 1 2, ix—.95 (AMNH). Black Mts.,
2 6, v.17— (AMNH). Madison County. Hot Springs, 1 2 (MCZ). Polk
County. Columbus, 1, vi.19.52; 1, vii.14-21.52 (USNM). Guilford County.
Greensboro, 1, viii.9.—, F. C. Pratt (USNM). OHIO. Athens County. Athens,
2, x.9-15.49; 3, vi.22-30.50. Waterloo Tnsp., 1, iv.23.50; 2, vi.27.50; 2, vii.6.50,
P. J. Spangler, Jr. (USNM). Hamilton County. Cincinnati, 1, viii.17.—, H.
Soltau (USNM). Richland County. Mansfield, 1 2 (ORES). Ross County.
Londonderry, 2, vili.5.50, P. J. Spangler, Jr. (USNM). Scioto County. Near
Friendship, Roosevelt Lake, 3, vii.29.50, P. J. Spangler, Jr. (USNM). Vinton
County. Lake Hope, 2 6, 1 2, vii.22.52, W. C. Stehr (CAS); 1, x.7.58, W. C.
Stehr (NL). Warren County. Mason, 2 m. S., 12 6, 7 2, vi.24.57, J. R.
Zimmerman (NMSU). OKLAHOMA. Atoka County. Atoka, 1 2, vi.13-15.—,
Wickham & Fenyes (CAS); 1, vi.13-15.—, Wickham (USNM). Bryan County.
Lake Texoma State Park, 2 ¢, 2 2, vi.24.62, E. Van Tassel (CNA). Marshall
County. 1, vi.5.53; 1, viii.5.53 (USNM). Ottawa County. 1 4, vi.5.31,
Costner & Davis (MCHS). Payne County. Stillwater, 1é , 1 9, ii.10.38,
K. C. Emerson (MCHS). Pittsburg County. McAlester, 1 6, vi.11.—, Wick-
ham & Fenyes (CAS); 3, vi.11.—, Wickham (USNM). PENNSYLVANIA.
Adams County . Arendtsville, 1, viii.4.51, P. J. Spangler, Jr. (USNM). Starrs
Mill, 1, vi.15.49, P. J. Spangler, Jr. (USNM). Allegheny County. Pittsburgh,
36,42; 16, vi.l4—; 2 29, vii——; 1 2, viii——; 1 9, ix.2.—; 1 Q,
ix.4.— (CNG). County, 4, iv.5.90, E. A. Klages (CNL); 3 6, 8 2 (CNG).
Delaware County. 1 6, v.18.— (ANSP). Franklin County. Chambersburg,
1, x.1.—; 2, J. D. Sherman, Jr. (USNM). Juniata County (?). Path (Val-
ley?), 8, J. D. Sherman, Jr. (USNM). Lebanon County. Lebanon, 10, J. D.
Sherman, Jr. (USNM). Philadelphia County. Philadelphia, 2 (USNM).
Philadelphia? Neck, 1 6, 2 2, v.12.—, Blaisdell (CAS). Phila. Neck, 2 é,
Laurent (AMNH);2 ¢,7 2, v.4.—; 1 6, 4 9, ix.13.— (ANSP); 1; 1, vii.15.04;
1, v.2.—, Kaeber (USNM); 2, vi.9.00; 1, ix.13.—, G. M. Greene; 2, P. Lauren
(USNM). Westmoreland County. Jeannette, 1 ¢,5 2, vii——:; 1 4, vii.3.—;
OP vit —— 6) Oe avin Oe vir te de 1 Or vi24 ——- 2S. 3°
vi.27.— (CNG). SOUTH CAROLINA. Beaufort County. Beaufort, 4 ¢,
3 2;2 6, iv.12.92: 1 6, iv.18.92, G. D. Bradford (AMNH). Kershaw County.
Camden, 4 6, 1 2, vi.23.29, P. J. Darlington, Jr. (MCZ). Pickens County.
Clemson, 2, xi.25.27, D. Dunavan (USNM). TENNESSEE. Davidson County.
Nashville, 1, viii.4-15.97, Wickham (USNM);: 1, Osburn (USNM). Fentress
County. Aijlardt, 1 4, viii.17.22, T. H. Hubbell (MCHSU). McMinn County.
Athens, 1 2, vi.24.63, A. R. Moldenke & J. A. Woods (AMNH). Shelby
County. Memphis, 1 6, v.16.12, Blaisdell (CAS). TEXAS. Brazos County.
College Station, 1, iii.31.35, H. J. Reinhard; 1, vii.19.59, H. R. Burke (TAM).
Collin County. Plano, 1, vii—.07, E. S. Tucker (USNM). Harris County.

MEM. AMER. ENT. SOC., 26

80 THE GENUS LACCOPHILUS

Houston, 5, vi.15.32, Harwood (CNL). Harrison County. Karnack, 4 6,7 2,
vi.17.58, R. L. Fischer, J. Eichmeier & R. A. Schneibner (MCHS). Panola
County. Carthage, 1 6, v.8.52, M. Cazier, W. Gertsch & R. Schrammel
(AMNH). VIRGINIA. Albemarle County. Ivy (Depot?), 5 2, vi.—.33,
F. E. Winters (CAS). Alexander County. 1, vi.18.10; 1, vii.7.08 (USNM).
Bath County. Warm Springs, 1 2, x.1—, Leng & Van Dyke (CAS). Fairfax
County. Great Falls, 2, i.9.06; 1, iii.17.07, D. H. Clemons (USNM). County, 2,
ix.20-22.27, Shoemaker (USNM). Henrico County. Richmond, 4 é, Slossom
(AMNH);1 29 (MCZ). Nansemond County. Suffolk, 10 ¢, 6 2, vi.27.1876,
Dimwock (MCZ). Nelson County. 5, viii.6.16, W. Robinson (USNM).
Nottoway County. Camp Pickett, 2 6, 1 2, v.1.52, B. Adelson (CAS). Spotsyl-
vania County. Fredericksburg, 1, ii1.16.89; 3, iii.15.91; 2, x.4.97; 3, i11.19-26.99,
W. D. Richardson (USNM); 3, ix.—.98, J. D. Sherman, Jr. (USNM); 1 4,
12, i.—.22 (AMNH). VERMONT. Bennington County. County, 1 6, 1 9,
vil.—.94, C. H. Roberts (AMNH);1 2 (ASNP). WEST VIRGINIA. Green-
briar County. White Sulphur Springs, 1 °, vii—. —, A. Fenyes (CAS); 10,
vili.23.10, W. Robinson (USNM). Jackson County. Ripley, 3, vi.25.30, F. N.
Musgrave (CNL). Jefferson County. WHarper’s Ferry, 4, ix.12.17, J. E. Bene-
dict, Jr. (USNM). Kanawha County. County, 2 é, 3 92, vii.—.35 (CNG).

Laccophilus fasciatus terminalis Sharp, new status
(Figs. 46, 48, 53, 282, 283)

Laccophilus terminalis Sharp, 1882a, p. 292. Type: British Museum (Natural
History), female, Texas; type locality here restricted to Davis Mountains,
Jeff Davis County, Texas; Sharp, 1882b, p. 11; Horn, 1883, p. 277; Horn,
1894, p. 313; Leng, 1920, p. 77; Zimmermann, 1920, p. 26; Leech, 1948b,
p. 402-3; Leech and Chandler, 1956, p. 314; Zimmerman and Severin,
1957, p. 30.

DIAGNOSIS. — The elytral pattern serves as the best recognition character
for L. f. terminalis. The irrorated dots are sparsely applied, and the pattern
outline is distinctly margined with dark brown or black. On the posterior half,
there is a tendency to form a fascia or blotch similar to rufus and fasciatus.
It is never complete, however, and, in some specimens, the pigment is restricted
to patches just behind the mid-lateral spots. Megascopically terminalis often
has a checkered appearance. Small individuals of L. sonorensis resemble termi-
nalis, but they are not irrorated. The male lacks a file, and the female has
rakelike ovipositor. Some specimens of mexicanus and salvini have a tendency
to form a dark blotch on the posterior half of the elytra, but are black beneath
(except female mexicanus atristernalis in coastal California and Oregon).

NOMENCLATURAL NOTES. — There appears to be no syno-
nymical problems in this race, but it is reduced to subspecies since it
intergrades with fasciatus and rufus.

VARIATION. — This race poses one problem in that the elytral
pattern can vary markedly from the usual pattern. There is a gen-

JAMES R. ZIMMERMAN 81

eral fading that causes some difficulty in recognition. The basic
ground pattern remains, however. The fading is most common in
individuals from the southwestern United States.

More population data was accumulated for this race than any
other in the study. Size and the PW/EL ratio is the same for males
and females. The average size (4.8 mm) is larger than for fasciatus
and most rufus populations (Table 2). There is considerable geo-
graphic variation in size. Central and south Texas populations are
about the same as midwestern rufus, but are smaller than other ter-
minalis to the west and south. Populations from northern New Mexico
south to Durango, Mexico, and from west Texas to California appear
to be much the same. South and east of Durango, however, there is
an increase in size. From Linares, Nuevo Leon, to Jalisco, the popu-
lations have an average length of almost 4.92 mm which is 0.05 to
0.10 mm larger than in other parts of the range. One exception is a
single large sample from about the middle of the Baja California pen-
insula which is equally long (4.95 mm). Farther south around La
Paz, the value is about the same as that found from New Mexico to
California.

It seems that while terminalis retains a fairly uniform pattern (ex-
cept for fading) and shape throughout its range, it is still undergoing
considerable change in size from one region to another. This should
not be surprising, since it covers an area with so much physiographic
contrast and climatic variation. About the only uniform feature in
its distribution is the preference for an aquatic habitat in an arid en-
vironment. It is perhaps more difficult to understand why other Lac-
cophilus which cover equally diverse areas (for example, L. maculosus
decipiens and L. maculosus shermani) do not show this degree of
variation.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— L. f. terminalis occurs from southwestern South Dakota to Jalisco.
Its eastern boundary approximates the ninety-seventh parallel where
it is replaced by rufus. From South Dakota the limits swing sharply
south and westward to the Rocky Mountains; and its northwestern
boundary is through central New Mexico, Arizona, and southern Cal-
ifornia. Outlying localities are known from North Dakota, northern
and southwestern Utah, Marin County of California, and southwest
Oregon. Numerous other collections in the intervening areas have

MEM. AMER. ENT. SOC., 26

82 THE GENUS LACCOPHILUS

not yielded additional records, however. It apparently occurs through-
out Baja California, and its southern limits are fairly well defined by
the Mexican Plateau and the central Mexican Highlands.

It is probably the most common Laccophilus (and any other dytis-
cid) in the southwestern United States and northern Mexico. It occurs
in practically any small aquatic habitat and, apparently, migrates very
readily since it often appears in street gutter pools within a few days
after a rain. L. f. terminalis is usually in heavier clay or adobe soils
that are exposed to bright sun and have considerable debris; it can
also be found in small, sluggish, streams. It ranges from sea level to
about 6000 feet, but may occasionally be taken above that elevation.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Benson,
2m. E., 2 6,8 2, x.6.63; 4 m. W., 4 2, x.6.63, K. L. McWilliams (NMSU);
5 6, 2 9, ii.25.33, O. Bryant (CAS). Douglas, 2 ¢, 1 9, iii.15.33, W. W.
Jones (BERK). Fairbank, S. Pedro R., 6, iii.12.36, J. G. Needham (CNL).
Huachucha City, 8 6, 17 9, ix.6.61, JRZ (NMSU). MHuachucha Mtns., 13,
v.7-8.53, A. & H. Dietrich (CNL); 4 ¢, 4 2, Barber (AMNH). Portal, 5 m.
W., SWRS., 1 6, vi.20.55, M. Statham; 2 ¢, x.1.55, M. Cazier; 9 6, 5 9,
x.21.55, E. Ordway; 1 9, iv.3.56; 1 6, iv.12.56; 1 2, iv.21.56, E. Ordway;
1 6, v.4.56, M. Statham (AMNH); 2 6, 2 9, x.21.55, E. Ordway (UMMZ)
comp. with type in BM(NH) by FNY; 5 m. SWRS., 8 6, 12 2, —.——,
L. B. Bartholomew (NMSU); 2 m. NW. Rodeo, New Mexico, 3 ¢, ix.5.61,
JRZ (NMSU). St. David, 2 6, 2 2, vii—.44, G. E. Pickford (FM); 2 ¢é,
2 2, vii.15.44, G. E. Pickford (CAS). Tombstone, 5m. N., 3 6, 8 2, ix.6.61,
JRZ; Willcox, 11 m. W., 5 6, 1 2, x.6.63, K. L. McWilliams (NMSU). Gila
County. Globe, Pinal Ck., 75, iv.25.53, A. & H. Dietrich (CNL); 1 4, 1 9,
x.13.48, F. H. Parker (CAS). Rice (San Carlos R. Res., R. R. Sta.), 4,
vii.9.31, D. Parker (CNL). Graham County. Near Artesia, rd. to Marijilda
Canyon from Hwy. 666, 4 4, 1 92, vii.3.65, H. B. Leech (CAS). Pima, 2,
x.21.24, A. A. Nichol (ARI). Pinalena Mtns., Hospital Flat, 1 2, viii.3.65,
H. B. Leech (CAS). Maricopa County. Gila Bend, 1, xii.13.45, H. P. Chan-
dier (BERK). Phoenix, 1, vi.23.94, J. D. Sherman, Jr. (USNM); 44 ¢, 51 2,
Kunze (AMNH); 1 8, v.31.40, G. P. MacKenzie (AMNH). Wickenburg,
2 6,5 9, iv.16.62, JIRZ (NMSU); 5 m. E. Hassayampa R., 1 6, 1 9, iv.2.66,
J. Schuh (SCH). Pima County. Ajo Mtn., Alamo Can. (OPCNM), xii.19.39,
C. F. Harbison (BERK). Ajo, 2 9, viii.31.55, P. S. Bartholomew (CAS).
Catalina Mtns., 9, iv.18.—, Hubbard & Schwarz (USNM); 1, H. & S. (USNM);
Lowell Ranger Sta., 1, vii.6-20.16, J. D. Sherman (USNM); 2, x.21.24, A. A.
Nichol (ARI); Bear Can., Molino Basin, 2, iv.18.—, v.20.53, A. H. Dietrich
(CNL). Florida, 1, xi.18.39, P. Lightle (ARI). Madera Can., 1 ¢, 1, ix.2.54,
Menke & Stange (DAV). Pantano, Cienega Ck., 6, iii.25.53, A. H. Dietrich
(CNL). Quitobaquito, OPC, Ntl. Mon., 12, iv.11.53; 2, vi—53, A. & H.

JAMES R. ZIMMERMAN 83

Dietrich (CNL). Santa Rita Mtns., Florida Can., 3, iv.19.59, F. G. Werner
(ARI); 2 6, 4 9, ix.7.61; 7 6, 7 92, xii.2.61; 1 9, viii.6.62, JRZ (NMSU).
Sta. Rita Mtns., 1 2, vii.25.41, O. Bryant (CAS). South Catalina Mtns., 2 6,
1 6, iv.10.38, F. H. Parker (DAV). Sta. Catalina Mtns., 2 2, x.21.24, A. A.
Nichol; 1 6, 1 9, v.8.33, O. Bryant; 2 4, xi.20.33, O. Bryant; 1 3, 6 Q, i.2.38,
Van Dyke; 1 6, 1 2, v.16.39, E. P. Van Duzee (CAS). Tanque Ck., 1,
ili.14.36, J. G. Needham; 8, iii.28.53, A. & H. Dietrich (CNL). Tanque Verde,
32 6, 42 Q, vii.25.61, JRZ; Tucson, 15 m. NE., 1 6, x.5.63, K. L. McWilliams
(NMSU); 10 6,5 9, vi.1.33, O. Bryant; 4 6, iii.22.33, O. Bryant; 3, iv.25.—,
Hubbard & Schwarz (USNM); 1 ¢, iv.15.33, O. Bryant (CAS); 20 m. E., 3 4,
1 9, ii.16.66, J. D. Vertree (SCH); 1 6, 2 9, iv.5.97, Koebele (CAS); 2 2,
ix.—.28, F. H. Parker (CAS); Sabino Can., 1, x.31.18, A. G. Boring (USNM).
Pinal County. Aravaipa Can. (Near Mammoth), 1, v.21.53, A. & H. Dietrich
(CNL). Picacho, 2 ¢, 3 9, iv.19.33, O. Bryant (CAS). Santa Cruz County.
Nogales, 1 6, 1 2, iv—.97, Koebele (?) (CAS); 5 m. N., 13 6, 9 Q, ix.6.61,
JRZ (NMSU). Patagonia, 1, v.26.53, A. & H. Dietrich (CNL); 1 ¢, 1 2,
ix.17.52, B. Malkin (FM); 3 m. S., Sonoita Ck., 1 é, 1 2, ix.6.61, JRZ
(NMSU). Pena Blanca L., Coronado Ntl. For., 6 ¢, ix.6.61, JRZ (NMSU);
1, iv.4.66, J. Schuh (SCH); Pajarito Mtns., 3 6, 1 9, viil.61; 14, 2 9,
viii.14.61; 14 6,9 9, vi.28.62; 1 @, vii.3.62; 3 6, 1 2, vii.6.62; 1 3, vii.12.62;
14 $, 10 @, vii.19.62; 1 6, 2 Q, vii.20.62; 1 3, vii.24.62; 1 6, vii.25.62; 1 9,
vili.8.62; Arnett & Van Tassel (CUA). Yavapai County. Bloody Basin, 2 ¢,
vi.8.47, F. H. Parker (CAS). Castle Hot Springs, 3 2, iv.5.42 (AMNH).
Minnehaha, 38 (USNM); 11 ¢, 18 2, Barber (AMNH). Prescott, 7, J. D.
Sherman, Jr. (USNM); 5 é, 10 2, vii—.—, Hulst (AMNH); 1 2 (ANSP).
Yuma County. County, 2 2, vi—— (LACM). CALIFORNIA. Los An-
geles County. Long Beach, 3 6, 6 2, viii—.11 (FM). Imperial County.
Brawley, 3, vili.8.14, J. C. Bradley (CNL). Desert Shores, 2 m. S., 1 2,
iv.24.62, JRZ (NMSU). Imperial, 1, iv.24.25 (BERK); 1 6, 1 2, same data
(CAS). Imperial County, 2 6, 6 9, v.25.25, J. J. Gehring (?) (MCZ); 2 é,
4 2, iv.3.24 (CNG). Inyo County, China Ranch, SE. Tecopa, 2 4, iii.23.67,
H. B. Leech (CAS). Marin County. Marin County, 1, x.18.19, H. Dietrich
(CNL). Mono County. Mammoth, 3 6,5 2, viii.17.52, PSB (CAS). Orange
County. Laguna Can., 3 9, ii.14.59, M. E. Irwin (DAV). Corona del Mar,
1 6,1 9, xii.27.58, M. E. Irwin (DAV). Lower San Juan Cpgd., 1 6, xi.6.54,
Menke & Strange (DAV). Riverside County. Blythe, 3 6, 2 92, iv.17.62, JRZ
(NMSU). Camp Pendleton, 1, x.23.45 (BERK). Riverside, 7, F. E. Winters
(CNL). San Bernardino County. Earp, 10 m. NE., 4 é,7 2, viii.16.63, C. D.
MacNeill (CAS). San Diego County. Borrego, 1 6, 1 2, x.28.39, G. P.
Mackenzie (LACM). Borrego Val., Frenchy’s Well, 1 2, i.2.35, H. J. Rayner
(CAS). Descanso, 1 4, ix.23.52, B. Malkin (FM). Escondido 12 m. N., 5,
ii.20.55, 1, ix.21.54, R. K. Benjamin (USNM); 10 m. N., 1, xii.17.54, Scissors
Crossing, San Felipe Ck., 1 ¢, 1 2, iv.15.65, H. B. Leech (CAS). Warner’s
(Springs?), 2 6, 4 2, vi.18.25, J. G. Gehring? (MCZ). Yuba County. Marys-
Ville, viii.1.34, R. Wagner (CAS). COLORADO. El Paso County. Nr. Palmer

MEM. AMER. ENT. SOC., 26

84 THE GENUS LACCOPHILUS

Lake, 1 9, vili.15.61, JRZ (NMSU). Kiowa County. Eads, 8 m. S., 4 6,
1 9, ix.11.63, K. L. McWilliams (NMSU). Lincoln County. Hugo, 8 m. SE.,
1 6, ix.11.63, K. L. McWilliams (NMSU). Sedgwick County. Ovid, 1 6,1 9,
vill.17.61, JRZ (NMSU). KANSAS. Gray County. Cimarron, 3 é, 1 @,
vili.22.56, P. J. Spangler (USNM). Hamilton County. Syracuse, 7 m. E.,
2 6, 1 9, ix.11.63, K. L. McWilliams (NMSU). Seward County. Liberal,
48,5 9, vili.12.58; 3 m. N., 2 6, 3 @, viii.12.58; Near Kismet, Cimarron R.,
1 2, vii.12.58, JRZ (NMSU). NEBRASKA. Lincoln County. North Platte,
10 m. W., 1 Q, viii.17.61, JRZ (NMSU). NEW MEXICO. Colfax County.
Cimarron, 10 m. N., 1 6, vii.15.65, A. H. Smith; Near Maxwell, 1 ?, viii.14.61,
JRZ; Raton, 1 9, vii.5.63, R. N. Gennaro; 7 m. E., 1 ¢, vii.15.65, A. H. Smith
(NMSU). Dona Ana County. Hatch, 1 6, 1 2, xii.14.45, H.P. Chandler
(FM); 7 m. W., 4 6, 4 2, iv.20.62; Jornado del Muerto, 15-20 m. N. of Las
Cruces, Taylor Wells, 3 6, 4 2, vii.28.63; Jornado Range, 1 6, 4 2, vii.20.63;
Ropes Springs, 3 6, 11 2, xi.18.63, K. L. McWilliams; Las Cruces, 6 6, 2 2,
vii.5.61; 2 4, vii.9.61; 3 2, vii.10.61, JRZ; 14 4, 12 9, vii.6.61, E. E. Staffeldt
& JRZ; 1 6, x.1.62, K. L. McWilliams (NMSU); 10 m. E., Organ Mtns., 5 é,
3 9, vili.7.65, H. B. Leech (CAS). Mesilla, 1 m. SW., 3 6, 3 2, vii.6.61;
1 6,2 Q, vil.13.61;6 6,5 Q, vii.13.61, JIRZ (NMSU). Eddy County. Queens
(El Paso Gap), 2, xi.9.14, E. G. Holt (USNM). Grant County. Hurley,
13 6, 2 9, vii.17.61; Lordsburg, 13 m. E., 3 6, 2 2, x.6.63, JRZ; Silver City,
10 m. W., 1 6, 2 9, vii.16.63, R. D. Ohmart (NMSU). Guadalupe County.
Santa Rosa, 1 6, 1 92, viii.24.57, D. Lauck (USNM); 20 m. SE., 12 6, 13 Q,
vii.29.63, R. D. Ohmart; 5 m. NE., 3 6, 5 9, ix.10.63, K. L. McWilliams
(NMSU). Harding County. Mosquero, 10 m. E., 32 6, 25 2, x.23.65, A. H.
Smith (NMSU). Hidalgo County. Animas, 5 m. S., 16 6, 11 2, vii.24.61,
JRZ; 12 m. SE., 115 6, 91 9, xi.23.62, K. L. McWilliams (NMSU); 8 m. S.,
2 9, vii.31.65, H. B. Leech; 15 m. S., 1 92, vii.31.65, H. B. Leech (CAS).
Cienega Lake (Nr. Rodeo), 2 2, vii.17.64, R. H. Arnett, Jr. & E. Van Tassel
(CUA). Double Adobe Rnch., Animas Mts., 5500 feet, 2 6, 3 9, viii.15.52,
H. B. Leech (CAS). Gary, 5 m. W., 7 6, 2 2, vii.24.61, JRZ; Lordsburg,
12 m. W., 4 2, x.6.61, K. L. McWilliams; Nr. Cloverdale, 6 ¢, 6 9, vii.24.61;
Roadforks, 2 6, 5 2, ix.5.61; Rodeo, 12 m. N., 18 6, 7 2, ix.5.61, JRZ
(NMSU). Lincoln County. Alto, El. 7000 feet, 7 ¢, 10 2, vil.27.61, JRZ;
Carrizozo, 13 m. NE., 2 6,5 9, ix.10.63, K. L. McWilliams (NMSU). Angus,
Rio Bonito, 1 4, viii.7.65, H. B. Leech (CAS). Luna County. Columbus,
1m.S., 11 ¢, 12 9, vii.17.61, IRZ (NMSU). Deming, 15 m. S., 2 6, 4 2,
vii.17.61; 15 m. NW., 4 4, 2 9, vii.17.61, JRZ (NMSU); 16 m. W., 2 ¢, 3 @,
x.6.63, K. L. McWilliams (NMSU). Mora County. Mora, 10 m. S., 4 6,
8 2, v.1.66, A. H. Smith (NMSU). Otero County. Alamogordo, 1 6, V.
Krockow (AMNH). Pinon, 40 m. S., 10 6, 2 2, vii.29.63, R. D. Ohmart
(NMSU). Quay County. San Jon, 1 6, 1 9, 1x.10.63, K. L. McWilliams
(NMSU). Tucumcari, 10 m. W., 1 @, ix.10.63, K. L. McWilliams (NMSU).
Rio Arriba County. Canjilon, 3 m. W., 1 6, 2 2, ix.25.65, A. H. Smith; El
Rito, 1 4, vi.23.65; Ojo Caliente, 1 2, vi.23.65, A. H. Smith, R. L. Smith, &
JRZ; Velarde, 4 6, 1 9, vi.22.65, A. H. Smith (NMSU). Roosevelt County.

JAMES R. ZIMMERMAN 85

Causey, 1 6, 3 2, viii.3.65, J. E. Sublette (ENMU). San Miguel County.
Las Vegas, 1, ii.2.02 (CNL). Mosquero, 19-20 m. W., 37 6, 31 2, x.23.65;
Trujillo, 19 m. E., Tremintina Ck., 39 2, 36 2, x.23.65, A. H. Smith (NMSU).
Sierra County. Hillsboro, 4 m. E. (label says Pedra R. Gorge, but must mean
Percha Ck.), 3 6, 2 9, viii.31.52, B. Malkin (FM). Kingston, 5 m. E., 1 2,
v.27.66, JRZ (NMSU). Socorro County. Nr. Bingham, 1 2, vii.21.62, R. D.
Ohmart (NMSU). Taos County. Taos, 4 m. E., 2 2, vii.14.64, A. H. Smith
(NMSU). Torrance County. Corona, 5 m. NE., 1 6, 3 2, ix.10.63, K. L.
McWilliams (NMSU). Union County. Pasamonte, 8 m. E. & 2 m.N., 2 46,
1 2, vii.15S.65, A. H. Smith (NMSU). NORTH DAKOTA. Barnes County.
Valley City, 1, ii.—.18, Gabrielson (USNM). OKLAHOMA. Cleveland
County. Norman, 1 2, iv.18.19 (CAS). Grady County. Chickasha, 1, vii.15.15,
W. H. Larrimer (USNM). Texas County. Guymon, 3 m. NE., 4 4, 4 2,
ix.11.63, K. L. McWilliams (NMSU). OREGON. Jackson County. Rogue
River, 2, ix.—.02, C. R. Biederman (USNM). SOUTH DAKOTA. Penning-
ton County. Hill City, 1 2, viii.5.35, A. Thrupp (NMSU). TEXAS. Bell
County. Ft. Hood, 5, vii.19.55, T. Matthews (CNL). Bexar County. Ft. Sam
Houston, 1 4, vii.13.52, B. J. Adelson (LACM): 1 6, vi—.52, B. J. Adelson
(CAS). Leon Ck., 10 6, 3 2, x.11-12.52, B. J. Adelson (CAS). San An-
tonio, 2 6, 1 2, Wickham (CNG). Blanco County. Cypress Mill, 2, ix.10.88;
5S (USNM); 1 6, 1 2, same date (CAS). Round Mtn., 2, Kaeber (USNM);
3 2 (MCZ). Shovel Mtn., 6 6,5 2, F. G. Schaupp (AMNH); 9 (USNM).
Brewster County. Alpine, 20 m. S., 2 2, v.12.27, J. D. Martin (CAS); 1,
vii.20-22.— (USNM); 1, vii.1-15.26, B. C. Pooling (CNL); 3 2 (MCHS).
B.B.N.P., Chisos Mtns., 1 4, vii.3.42, H. A. Scullen (ORES); 4 6,1 2, ix.5-6.52,
B. Malkin (FM); 7 m. W., Chisos Junct., 5 6, 5 2, viii.3.61, JRZ (NMSU).
Boquillas, 1 2, vii.7.48, C. & P. Vaurie (AMNH); 7 6, 7 9, viii.2.61, JRZ
(NMSU). Chisos Mtns., 1, x.12.08, Mitchell & Cushman (USNM). Mara-
thon, 10 m. E., 5 6, 12 9, viii.1.61, JRZ (NMSU); 6 ¢, 6 92, ix.12.49, O.
Bryant (CAS). County, 1 6, 1 2, v.3.27, J. O. Martin (CAS). Burnett
County. 8, Hubbard & Schwarz (USNM). Cameron County. Brownsville, 1,
iv.13.95, Townsend (USNM); 7, iii.5.04, H. S. Barber (USNM); 2, vii.m—-;
2 (USNM); 1 6, Wickham (ORES); 3 6, 3 2, Wickham (MCZ). Esperanza
Ranch, 4, viii.19— (USNM). Port Isabel, 1 ¢, 2 2, x.20.49, O. Bryant
(CAS). Culberson County. Nickel Ck. Sta., 2.5 m. E., 21 6, 22 9, 1x.2.52,
B. Malkin (FM). Van Horn, 1 6, vii.10.50, R. F. Smith; 1 6, vii.10.48,
C. & P. Vaurie (AMNH). Edwards County. Camp Wood, 2 4, 2 9, iii.8.33
(TAM). El Paso County. Clint, 1 m. SW., 17 6,6 9, vii.19.61, JRZ (NMSU).
El Paso, 9, vii.28.14, J. C. Bradley (CNL); 3 6, 1 2 (AMNH); 1 6, 2 9
(ANSP). Falls County. Reagan (Wells?), 5, iii.6.36, J. G. Needham (CNL).
Goliad County. County, 1, viii.17.—, J. D. Mitchell (USNM). Gonzales
County. Harwood, 1, Wickham (USNM). 4Harris County. Houston, 1,
Wickham (CNL). Hays County. Dripping Springs, 1 4, viii.9.42, W. S. &
E. S. Ross (CAS). Hidalgo County. Edinburg, 7, ii.24.36, J. G. Needham
(CNL). Hudspeth County. Dell City, 9 m. SW., 1 6, vil.31.50, R. F. Smith

MEM. AMER. ENT. SOC., 26

86 THE GENUS LACCOPHILUS

(AMNH). Ft. Hancock, 5 ¢, 3 9, vii.19.61, JRZ (NMSU). J. Davis County.
Davis Mountains, Limpia Cn. Ck., 3 6, 6 9, iv.19.53, B. Adelson & M. Wash-
bauer (BERK); 2 ¢, 8 9, v.9.51, O. Bryant; 1 6, vi.26.46, Van Dyke (CAS);
1 6, 1 9, vili.3.61; Madera Cn. Ck., 1 92, viii.3.61; 10 6, 2 2, x.27.61, JRZ
(NMSU); Elbow Cn. Ck., 7 ¢, 12 2, x.27.61; Phantom Lake, 2, vi.20.16,
F. M. Gaige (USNM). Jim Wells County. Alice, 8 m. NE., 25, viii.25.35,
C. E. Burt (USNM). Llano County. Near Llano, 5 6, 5 2, iv.25.63, G. Child
(NMSU). Pecos County. Pecos R., Rt. 290, 3 6, 4 2, iv.23.63; Sheffield,
1 2, iv.23.63, G. Child (NMSU). Presidio County. Marfa, 14 m. N., 3 ¢,
8 2, vil.1.61, JRZ (NMSU). Sutton County. Devil’s River (Nr. Sonora), 8,
v.3-6.07, Pratt & Schwarz (USNM). Travis County. Austin, 4 6,1 92, xii.5.28,
J.O. Martin (CAS). Uvalde County. Sabinal, 1, iii—.10, F. C. Pratt (USNM).
Val Verde County. Del Rio, 5, vii.23-24.—, Wickham (USNM); 7 4, 13 9,
x1.15.49, O. Bryant (CAS). Webb County. Laredo, 1 2, v.20-24.48, F. G.
Wagner & W. Nutting (ARI). Zapata County. Zapata, 12, ii.26.36, J. G.
Needham (CNL). UTAH. Ogden, 1 ¢ (Acc. 4858) (AMNH). Washington
County. Hurricane, 1 2, vi.22.59, R. D. Anderson. New Harmony, 1 2,
x.4.64; 28, x.11.64, R. D. Anderson (RDA). St. George, 2 6, xii.29.59,
G. Musser (CAS). WYOMING. Goshen County. Ft. Laramie, 3.5 m. S.,
2 6, vili.19.65, H. B. Leech (CAS).

MEXICO. — AGUASCALIENTES. Aguascalientes, 1 m.N.,4 6,1 2, vii.26.62,
JRZ (NMSU). BAJA CALIFORNIA. Agua Caliente (Cape district), 3 2,
iv.22.47, I. La Rivers (BERK). Arroyo de Calamajue, Near Calamajue, 1 ¢,
1 2, iv.9.61, A. G. Smith (CAS). Cabo San Lucas, 7.7 m. NE., 1 ¢, 4 9,
1.1.59, H. B. Leech (CAS). Catarina, 3, ix.2.51, G. A. Marsh; El Carrizal,
8 6, 7 9, iv.25.47, I. La Rivers (BERK). El Triunfo, 2 ¢, i.9.59; La Paz,
8.2 m. W. Hwy. Sur 1, 2 9, xii.31.58; 4 6,5 2, xii.31.58; 9.6 m. E., 1 6, 1 9,
x1i.30.58; 12.4 m. E., Las Cruces Road, 2 6, 6 2, xii.23.58, H. B. Leech; 4 92,
x.10.55, F. X. Williams; La Suerte, 1 ¢, 1 9, vi.4.63, R. K. Benjamin (CAS).
La Zapopita, Valle de Trinidad, 1 6, iv.10.61, F. S. Truxal (LACM). Las
Paras, 1, W. W. Mann (USNM). Miraflores, 3 m. NW., 1 ¢, H. B. Leech
(CAS). Padarone, Amarillo Arroyo, 12 6, 20 9, v.11.47, I. La Rivers (BERK).
San Antonio, 1 ¢, vii.12-17.19 (CAS). San Bartolo, 1 m. SE., 1 2, i.20.59,
H. B. Leech; 3 9, v.1.47, I. La Rivers (CAS). San Felipe, 2, J. D. Sherman,
Jr. (USNM). San Jose del Cabo, 4, Wickham (USNM). San Luis Gonzaga,
12 m. E., 45 6, 45 2, v.22.47, I. La Rivers (BERK). Todos Santos, 1 92,
iv.10.47, I. La Rivers; 9 m. S., 1 6, 2 2, 1.14.59, H. B. Leech (CAS). CHI-
HUAHUA. Camargo, 1 m. N., 15 ¢, 3 2, vii.25.62; Casas Grandes, 5 m. E.,
27 6, 26 2, xii.20.64, JRZ (NMSU). Catarinas, 5800 feet, 1 2, vii.25.47,
M. Cazier (AMNH). Chihuahua, 43 m. N., 143 ¢, 167 2, xii.8.62; Colonia
Juarez, 2 m. SW., 1 9, xii.20.64; Parral, 2 m. S., 1 2, vii.25.62; 10 m. S., 1 9,
xii.9.62, JRZ (NMSU). Primavera, 3 6, vi.30.47, M. Cazier, W. Gertsch,
R. Schrammel (AMNH). COAHUILA. Ramos Arizpe, 4 é6, 4 2, vii.7.63,
JRZ (NMSU). Villa Acuna, S. Lorenzo Rnch., 2 ¢, 3 2, vi.16.38, R. H.
Baker (TAM). COLIMA. Trapechi (near Colima), 1 6, 4 9, vii.30.62, JRZ

JAMES R. ZIMMERMAN 87

(NMSU). DURANGO. Abasolo, Rio Nazas, 3 6, 3 Q, vii.25.62;58 4, 61 2,
x.22.66, A. H. Smith & JRZ; Durango, 1 4, vii.26.62; 20 m. W., 1 4, xii.10.62;
San Juan del Rio, 10 m. N., 2 6, 4 2, xii.9.62, JRZ (NMSU). GUANA-
JUATO. Guanajuato, 2 6, 3 2, vi.20.57, D. R. Lauck (USNM). Irapuato,
5m.S., 10 6,4 2; Leon, 5m. S., 1 2, vii.26.62, JIRZ (NMSU). HIDALGO.
Zimapan, 4 6, 2 @, vii.9.63, JIRZ (NMSU). JALISCO. Ameca, 25 km. E.,
1 4, iii.25.64, JRZ (NMSU). Barranca Oblato, 1 2, x.23.66, A. H. Smith &
JRZ (NMSU). Cd. Guzman, 5 m. N., 2 6, 1 Q, ii.30.62, JRZ (NMSU).
Guadalajara, 8 m. S., 1 6, 2 2, ix—.54, F. X. Williams (CAS); 11 m. S.,
5 6,3 Q, vii.30.62; 20 m. E., 1 6,12, JRZ (NMSU). Jilquilpan, 20 m. W.,
1 6,5 9, xi.30.48, H. B. Leech, E. S. Ross (CAS). Lagos de Moreno, 2 ¢,
4 Q, vii.26.62, JRZ (NMSU). Mazamitla, 1 m. NW., 1 6, 1 9, it.8.53, I. J.
Cantral (UMMZ); 5 km. E., 1 3, 1 9, iii.27.64; Near Tala, 1 ¢, 2 9, iii.25.64;
Near Tizapan, Rd. 15, 13 6,9 9, iii.26.64, JRZ (NMSU). Ojuelos de Jalisco,
12m.S.,2 6, 3 9, xi.21.48, H. B. Leech (CAS). South of Guadalajara, 3 9,
vili.11.57, D. Lauck (USNM). Tecolotlan, 3 3, 4 2, x.24.66, JRZ (NMSU).
Tehetlican, Rd. 110, 1 6, 1 2, vii.28.62, JRZ (NMSU). Tlaquepaque, 1 °,
vii.—.53, N. L. H. Kraus (CAS). Union de Tula, 10 m. S., 2 6, 7 2, 11.28.64;
Zapotlenejo, 7 m. E., 4 6, 1 Q, iii.25.64, JRZ (NMSU). MICHOACAN.
Cuitzeo (near), 4 2, vii.27.62, JRZ (NMSU). Jacona, 3 9, vili.11.57, D. R.
Lauck (USNM). Jiquilpan, 8 km. E., 3 6, 2 2, JRZ (NMSU). Zamora,
9m. W., 1 6, xii.6.48, H. B. Leech (CAS); 1 6, 1 9, vii.28.62, JRZ (NMSU).
NAYARIT. San Blas, 5 m. E., 1 4, vii.31.62, JIRZ (NMSU). NUEVO LEON.
Arroyo de Lajillas, 10 m. S. Linares, 3 ¢, 3 2, xii.18.40, F. N. Young (UMMZ).
Linares, 20 m. W., Rio Linares, 6 6, x.16.48, H. B. Leech (CAS). Santa
Catarina, Huasteca Can., 5 6, 6 9, vii.7.63, JRZ (NMSU). PUEBLA. Near
Maria Andrea, 1 2, ix.10.64, JRZ (NMSU). QUERETARO. Queretaro,
17 6, 7 9, ili.27.63, IRZ (NMSU); 3 9, vi.20.57, D. R. Lauck (NMSU).
San Juan del Rio, 5 4, 1 9, iii.27.63, JRZ (NMSU). SAN LUIS POTOSI.
Agua Zarca (near Cd. del Maiz), Rd. 80, 3 6, 1 9, iii.25.63; El Salto, 1 2,
iii.25.63; Guaymuchil (near Naranjo), 1 ¢, iii.25.63; Jitalpa, 1 2, iii.23.63;
Presa de Guadalupe, Rt. 80, 5 6, 1 9, iii.26.63, JRZ (NMSU); Cuidad del
Maiz, 5 6,3 9, xi.20.48, H. B. Leech (CAS). Santa Maria del Rio, 4 6, 3 2,
1.26.63, JRZ (NMSU). SONORA. Aribabi (E. of Moctezuma), 4 ¢, 2 9,
xli.15.62; Bavispe, 1 6, 2 9, vii.26.63, JRZ (NMSU). Etchojoa, 7 m. NE.,
4 6, 10 9, 1.27.51, A. A. Hubert (CAS). Hermosillo, 1 2, ix.19-20.52, B.
Malkin & U. Thatcher (FM); 50 km. W., 2 6, viii.21.53; 1 92, vii.9-16.53,
B. Malkin (FM). ZACATECAS. Fresnillo, 16 m. NW., Rio Trujillo, 1 é,
vi.29.54, R. N. Brewer; Pinos, 10 m. S., 8 6, 5 9, viii.1.59, 15 m. S., 1 6,
vili.2.59, Ray Bandar (CAS).

INTERGRADATION IN LACCOPHILUS FASCIATUS

There are three zones of intergradation in Laccophilus fasciatus.
One is between L. f. rufus and L. f. terminalis, and two are between

MEM. AMER. ENT. SOC., 26

88 THE GENUS LACCOPHILUS

L. f. fasciatus and L. f. terminalis. Intergrades between rufus and
terminalis occur in a band that extends from central Texas to east
central Kansas at about the ninety-seventh parallel. Several hundred
specimens have been collected from several dozen localities in a strip
about 150 miles long that reaches from Dickenson County, Kansas,
to Noble County, Oklahoma (Tasch and Zimmerman 1961). In
that zone most of the specimens were more like rufus than terminalis
in elytral pattern and genitalia. Some intermediates were almost al-
ways present in any sample, however; and on a few occasions in
Harper, Kingman, and Sedgwick Counties, Kansas, the samples were
almost entirely of the terminalis pattern and genitalia. The overlap
in size and body proportions is great enough that it is not possible to
recognize clearly an intermediate condition in length or in the ratio of
pronotal width divided by length (Table 2). The mean values are,
perhaps, closer to the bulk of rufus size means than to those of ter-
minalis. Two large samples from Texas verify that the zone does con-
tinue southward below Oklahoma. In Texas there are also a sufficient
number of samples to illustrate clearly that the zone is a narrow one
that is probably less than 50 to 100 miles wide at the most.

Intergrades were not recognized soon enough in Kansas and Okla-
homa to find how far westward the zone extends. In the western
parts of those states, there is no evidence of intergradation. The lim-
its of the range of both races occur in South Dakota where they are
entirely allopatric (Zimmerman and Severin 1957). L. f. rufus is
confined to the extreme southeastern corner of the state and terminalis
to the southwestern one.

The situation between terminalis and fasciatus is more complex.
Intergrades occur in a long narrow strip from Pima county, Arizona,
along the west coast of Mexico to Nayarit and, perhaps, into Jalisco.
In Arizona most terminalis populations occur without any evidence
of intergradation with fasciatus, but the few specimens of fasciatus
that have been taken and previously identified as apicalis (its syno-
nym) show evidence of terminalis influence when closely examined.
Farther south along the coast, the general hybridizing of the two races
is more apparent in elytral pattern, genitalia, size, body proportions,
and larger average size of the males over females (Table 2).

In Jalisco there is evidence that the two races occur with no inter-
gradation. Not one intergrade has been found in more than 250
specimens from that state. In the usually diagnostic features of elytral

JAMES R. ZIMMERMAN 89

pattern, male genitalia, size, and body proportion, the populations
segregate into two distinct groups with no intermediates. One slight
bit of evidence for intergradation, however, is that a very large num-
ber of fasciatus from the vicinity of Autlan (74 males and 92 females)
have slightly larger mean values for length. The males are 4.80 mm
and the females are 4.74 mm which approaches the values for termi-
nalis populations. Usually the values are about 0.05 mm shorter in
both cases for fasciatus.

On the east coast of Mexico, there is also some slight evidence of
intergradation. In south Texas terminalis shows no influence from
fasciatus or rufus. A few (less than ten) specimens of fasciatus indi-
viduals with terminalis characters have been recognized, however.
Conversely, in southern Tamaulipas and northeastern tropical San
Luis Potosi, fasciatus generally occurs without evidence of terminalis
admixture; but occasional populations with intergrades do occur.

In all three instances, intergradation is clearly restricted to what
could be considered to be intermediate ecological areas in which inter-
grades might be as well adapted as parental genotypes. Between
rufus and terminalis the zone is in the ecotonal region between a
humid region and an arid one. In both instances of intergradation
between fasciatus and terminalis, the intergrades are between an arid
temperate climate and a humid tropical one.

The range of fasciatus approaches rufus in Texas, but as yet there
is no evidence of any contact between the two races. In some ways
these two races show more affinity than either does with terminalis.
They are more similar in elytral patterns, size, male genitalia, and
ecological preferences. It is possible that originally these two were
continuous while separated from terminalis; but, with the generally
increasing aridity of the Southwest and Mexican Plateau, terminalis
has increased its range more rapidly and has interposed between them
so that it now serves as the link between the races.

MATERIAL EXAMINED
L. f. fasciatus X L. f. terminalis

UNITED STATES OF AMERICA. — ARIZONA. Catalina Mtns., 2, xi.8.33,
Kath. Thomas (ARI); Bear Canyon, 3 é, 2 9, iii.26.46, J. W. Green (CAS).
Santa Rita Mtns., Florida Canyon, 1 ¢, 1 9, xii.2.61, JRZ (NMSU). Tucson,
1 $, ix.—.28, F. H. Parker (CAS). Santa Cruz County. Nogales, 1 6,

MEM. AMER. ENT. SOC., 26

90 THE GENUS LACCOPHILUS

iv—.97 (CAS). CALIFORNIA. Riverside County. Riverside, 1 ¢, F. E.
Winters (CAS). TEXAS. Kleberg County. Riviera, 1 ?, ii.28.36, J. G.
Needham (CNL). Not located. Eyle, 1 2, vi.15.57, David Lauck (USNM).

MEXICO. — NAYARIT. Acaponeta, 8 m. NW., Rio de las Canyas, 1 6,1 2,
xii.25.58, H. B. Leech (CAS). San Blas, 7 6, 11 2, ix.17-20.53, B. Malkin
(CAS); 1 @, ix.28.61, C. O. Morse (CAS); 5 m. E., 10 6, 8 Q, viii.31.62,
JRZ (NMSU). Sierra de Zapotan, 1 6, 3 2, xi—.42, Eugenio Paredes; Tepec,
18 m. NW., 4 2, xi.27.48; 20.5 m. NW., 1 6, xi.24.48, H. B. Leech (CAS).
SAN LUIS POTOSI. Agua Zarca (near Platanito), 4 4, 2 9, iii.25.63; Co-
moca, Rio Axtla, iii.23.63, JRZ (NMSU). C. del Maiz, 1 6, 1 2, xi.19.48,
H. B. Leech (CAS). Guaymuchil (near Naranjo), 1 6, 2 9, ili.25.63; El
Salto, 3 6, 10 9, iii.25.63, JRZ (NMSU). Jitalpa, 1 9, iii.23.63, JRZ (NMSU).
Palitla, 5 m. N. Tamazunchale, 4 ¢, 2 2, xii.22.48, H. B. Leech (CAS). Tama-
zunchale, 1 ¢, xii.30.47, Mulaik (CAS). Valles, 29 m. N., 1 6, viii.19.54,
F. N. Young (UMMZ). SINALOA. Concordia, 8 m. E., 1 6, 1 2, xii.12.62;
Culiacan, 10 m. N., 1 6, 8 2, viii.1.62, JRZ (NMSU). Elota, 6 6, 4 2,
vi1.27.56, R. & K. Dreisbach (MCHS); 1 m. S., 7 6, 7 9, viii.1.62, JRZ
(NMSU). Guasave, 4 6, 2 9, i.25.51, A. A. Hubert (CAS). Los Mochis,
6 6,6 8, vi.13.22, C. T. Dodds (CAS). Mazatlan, 7 6, 7 9, vili.1.62; 27 6,
292, xii.11.62; 7m. S., 9 6, 13 9, xii.11.62, JRZ (NMSU). Wolamo, 1 4,
vi.27.56, R. & K. Dreisbach (MCHS). (Villa) Union, 1 ¢, 1 9, vi.28.56,
R. & K. Dreisbach (MCHS). SONORA. Alamos, 1 9, 11.23.63, P. H. Ar-
naud, Jr. (CAS); 7 m. SE., 1 @, viii.12.60, P. H. Arnaud, E. S. Ross, D. C.
Rentz (CAS). Guaymas, 3, ix.29.23, W. M. Mann (USNM); 1 4, 1x.9.23,
W. M. Mann (CAS). Hermosillo, 50 km. W., Rancho San Francisco, 3 ¢,
vili.21.53, B. Malkin (CAS). Puerto Kino, 25 m. E., Rancho Montijo, 1 2,
vii.23.50, J. P. Figg-Hoblyn (CAS). TAMAULIPAS. Antiguo Morelos, 1 @,
11.23.63; 3 m. N., 1 6, 2 9, iii.26.63; Near El Limon, 1 é, iii.24.63; Llera,
Rio Guayalejo, 2 ¢, 8 9, iii.23.63; 20 m. S., 2 9, iii.24.63; Nuevo Morelos,
1 6, iii.25.63; Ocampo, 2 6, iii.24.63, JRZ (NMSU).

L. f. rufus X L. f. terminalis

UNITED STATES OF AMERICA. — KANSAS. Barber County. Isabel, 2,
iv.14.52, D. R. Bell (USNM). Dickenson County. Mt. Hope, 4 m. S., 7 ¢,
9 9, vil.8.60; 2 m. N., 3 6, 2 2, vii.15.60; Nr. Elmo, 11 6, 9 2, vii.15.60;
2 S$, vi.l1.58, JRZ (NMSU). Doniphan County. Wathena, 2 6, 1 9, C. T.
Brues (MCZ). Harper County. NE. corner of County, 7 ¢, 3 2, vii.20.60;
Danville, 2 m. S., 27 6, 14 9, vii.20.60, JRZ (NMSU). Harvey County.
Newton, 3 m. S., 1 4, vi.21.58; 2 m. SE., 1 6, 1 2, vi.21.58; 7 m. S., 3 é,
2 9, viii.12.59; 4 m. SE., 1 ¢, 10 Q, viii.12.59; 3 m. E., 12 6, 5 9, vii.6.60;
2m. E., 1 4, 3 9, vii.7.60; 4 6, 1 2, vii.6.60, JRZ (NMSU). Sedgwick,
5 $,6 Q, vil.13.60; Walton, 5 m. S., 1 9, viii.14.59, JRZ (NMSU). Kingman
County. Murdock, 2 m. S., 4 ¢, 3 2, vii.14.60; Waterloo, 5 m. E., 5 6, 2 92,
vii.14.60, JIRZ (NMSU). Marion County. Hillsboro, 4 m. NE., 1 2, viii.9.60;
6m. NE., 1 6, viii.9.60; Nr. Marion, 5 ¢, 7 9, viii.20.59; 2.5 m. NW., 1 ¢,
vili.9.60, JRZ (NMSU). Reno County. Haven, 8 m. E., 7 6,7 2, vii.13.60;

JAMES R. ZIMMERMAN 91

2 46,7 2, vii.19.60; 4 m. S., 4 6, 5 2, vii.19.60, JRZ (NMSU). Sedgwick
County. Cheney, 2 @, vi.5.58; Derby, 23 4, 28 2, vi.10.58; Kechi, 7 6, 10 2,
viii.10.59; Near Viola, 67 4, 422, viii.10.60; Near Bentley, 2 6, 4 2, iv.11.58;
St Mary's, 1 mi S:, 5 6, vit-1'3:60; Wichita, 3 m. E., 37 6, 35 9, ix.28°57;
74,7 8, iti.31.58; 5 9, iv.26.58; 3 6, 4 9, v.12.58; Wichita, 8 ¢, 2 2, vi.9.58,
JRZ (NMSU). Sumner County. WHunnewell, 14 6, 14 2, vi.22.60; 5 m. S.,
1 6, 1 2, vi.22.60; South Haven, 4 m. N., 1 4, vi.23.60; 3 2, vi.21.60, JRZ
(NMSU). OKLAHOMA. Kay County. Blackwell, 2 m. NW., 11 4, 10 2,
vi.20.60; 3 m. SW., 2 6, 3 92, vi.30.60; 3 m. E., 1 2, vi.30.60; Braman, 2 ¢,
4 2, vi.20.60: 8 m. E., 2 9, vi.20.60; 4 m. W., 1 9, vi.27.60; 2.5 m. NW.,
9 6,6 2, vi.28.60; 1.5 m. SW., 1 3, vi.28.60; Tonkawa, 1 m. S., 1 6, vi.23.60;
4m. NW., 18 6,7 @, vi.30.60, JRZ (NMSU). Noble County. Near Ceres,
3 6, 4 2, vi.23.60; Perry, 3 m. N., 8 6, 9 9, vii.1.60; 4 m. E., 7 6, 18 9,
viii.1.60, JRZ (NMSU). TEXAS. Dallas County. Dallas, 2 6, Wickham
(AMNH); 1 6, Leng, Van Dyke (CAS); 3, v.16.11, H. Pinkus; 1, Wickham
(USNM); 1 4, 1 2, Boll (MCHS); 1 ¢, 1 2, Boll (NMSU); 11 6, 19 2,
Boll (MCZ). Lee County. 1 (CNL); 6, J. D. Sherman, Jr. (USNM); 5
(USNM); Fedor, 1 ¢, v.26.09; 1 6, 1 9, vi—.— (MCZ).

Laccophilus proximus Say (Figs. 54-61, 289)

Laccophilus proximus Say, 1823, p. 101 (taken from J. L. LeConte’s editing of
the complete writings of Thomas Say, 1891, p. 514). Neotype: male,
Beaufort, Beaufort County, South Carolina, AMNH; Crotch, 1873, p. 400;
Schwarz, 1878, p. 438; Sharp, 1882a, p. 289; Blatchley, 1910, p. 210;
Blatchley, 1919, p. 308; Leng, 1920, p. 77; Zimmermann, 1920, p. 25;
Wilson, 1923b, p. 290; Young, 1953a, p. 34; Young, 1953b, Young, 1954,
p. 45; Zimmerman and Severin, 1957, p. 31; Zimmerman, 1960, p. 143.

Laccophilus americanus Aubé, 1838, p. 422; Sharp, 1882a, p. 291; Sharp,
1882b, p. 11.

Laccophilus confusus Sharp, 1882a, p. 292; Sharp, 1882b, p. 11.

DIAGNOSIS. — Recognition of this species is sometimes difficult without
examination of male genitalia. It belongs to the irrorated group with a coxal
file and sawlike ovipositor. It differs from L. maculosus in its small size (under
4.5 mm as compared to usually over 5.0 mm). It has less darkening and
coalescence of dots on the elytra than does L. m. maculosus or L. fasciatus;
and it is brown, yellow-brown or reddish-brown beneath unlike L. mexicanus or
L. salvini. Superficially its elytral pattern is similar to L. fuscipennis, L. vaca-
ensis, and L. spangleri, but differs as seen in figures 289, 290, 312-314. The
metacoxal file is weak in fuscipennis males; and there is usually an epipleural
flange in females of fuscipennis, but rarely in proximus. L. vacaensis and
L. spangleri lack a file and have rakelike ovipositors.

DESCRIPTION. — Small to medium (length 3.8 to 4.6 mm; width 2.1 to
2.4 mm), brown, irrorated species; metacoxal file prominent in males and weakly
present in females; prosternal process short; ovipositor sawlike. COLOR.

MEM. AMER. ENT. SOC., 26

92 THE GENUS LACCOPHILUS

Head: yellow above and beneath with strong reddish-brown tinge between the
eyes in the occipital region; appendages pale yellow except mandibles which
darken to brownish-red toward tip. Pronotum: about the same color as head,
but tending toward paler yellow at sides. Elytra: pale brownish-yellow back-
ground with irroration forming a pattern as shown in figure 289; coalescence of
pigment slight and rarely in specimens from north of Florida in the United
States, with no intensification of pigment at pattern margin as in maculosus;
epipleura pale yellow anteriorly, darkening to brownish-red in posterior con-
stricted portion. Tergite VIII: pale yellow or pale brown in the distal half,
varying from nearly black to pale brown in proximal half. Venter and geni-
talia: varying from brownish-yellow to brownish-red, seldom dark reddish-
brown as in maculosus. ANATOMY. Méicroreticulation: weakly double on
head, pronotum, and elytra; secondary mesh discernible, but individual cellules
still apparent. Head: supraclypeal seam closely parallel to margin. Pronotum:
WH/PW, 0.71; LP/PW, 0.40. Elytra: epipleura rarely with flange in United
States populations, but common in Mexico and West Indian specimens; apices
slightly truncate. Venter: coxal file prominent in males and composed of 24
to 28 lines; much weaker in female and composed of about 10 lines; prosternal
process with well-defined crest; lobes of postcoxal processes rounded and later-
ally projecting well posterior to midline; small protuberance on the middle of
the left posterior margin of the male fifth abdominal segment; hind margin of
male and female last ventral segment scarcely or not at all produced; small
asymmetrical crest on male segment, but smoothly rounded in female. Legs:
male pro- and mesotarsi enlarged in dorsoventral plane; palettes easily distin-
guished at 20 power magnification; fifth tarsal segment of proleg twice as long
as fourth, and fifth segment of middle leg one and two-thirds as long as fourth;
profemoral setae (6 to 7) shorter and finer than mesofemoral ones (5 to 6).
Genitalia: oval plate produced to acuminate tip with prominent ventral crest
curving strongly to the right; 8 to 10 raised lines on the right of the crest and
3 to 4 on the left; left paramere with an asymmetrical apex; right paramere
with produced apex; aedeagus evenly curved and tapered to apex; ovipositor
with about 13 sawlike teeth on each valve.

NOMENCLATURAL NOTES. — Young (1954) expressed the
opinion that proximus represents a number of allopatric subspecies.
Since americanus Aubé was described from the West Indies, it is
clearly allopatric to mainland populations, and it does appear to be
different. It is not clear what the nature of the variation is and how
the different populations are distributed throughout the Caribbean,
however, so no subspecific status for americanus is presented in this
paper. All mainland populations appear sufficiently similar to retain
in a single race, including Sharp’s confusus from Tabasco, Mexico.
Sharp thought that proximus was the same as maculosus, but these
are Clearly two different broadly sympatric species.

95

ZIMMERMAN

JAMES R.

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MEM. AMER. ENT. SOC., 26

94 THE GENUS LACCOPHILUS

VARIATION. — Elytral pattern is remarkably uniform in con-
tinental proximus. Specimens from southern Mexico, Kansas, and
Florida appear to be drawn from the same sample. Color differs in
specimens from southern Florida and from some of the western states,
however. Young (1954) says that proximus from southern Florida
are brighter than those to the north. Very pale individuals appear in
the populations as one moves across the plains states, but the per-
centage is never very high. Males and females show no difference in
size or shape and have been combined in Table 3.

Neither morphology nor quantitative data support retaining con-
fusus as a separate subspecies. It was described by Sharp from
Villahermosa, Tabasco, but samples from nearby states in southern
Mexico yield the same values as those from the United States. In
fact, there is greater variation among northern localities than between
Mexico and the United States. South Texas (Cameron Co.), Trans-
Pecos, Texas and Florida have mean values considerably lower than
other localities. They may represent real geographic differences; but,
unfortunately, sample size is comparatively small for all three and one
hesitates before concluding that these are different. Young had felt
that southern Florida populations were smaller than those from north-
ern Florida, however. Also, it would not be surprising if populations
on the edge of the range such as the Trans-Pecos one did differ from
the rest of the species.

There are anomalous values for the WP/EL ratio (Monroe Co.,
Indiana, St. Landry Co., Louisiana) which I am inclined to believe
are due to incorrect measurement of the elytral length. Other nearby
samples and other measurements do not support any real differences
in the populations; thus measuring errors for these two samples can
be suspected. The WP/EL ratio is about 0.555 for proximus, if the
two samples in question are not included.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This is one of the most widespread species in North America.
Even though it has a very large range, there appears to be little evi-
dence of racial differentiation on the continent. In the West Indies
there does seem to be a different race, however. The definition of its
range is beyond scope of this study, and only locality records for the
West Indies are included. L. proximus occurs from southeastern Can-
ada to Florida, westward to eastern Wyoming, Colorado, northeastern

JAMES R. ZIMMERMAN 95

New Mexico, all of Texas, and southward in eastern coastal Mexico
and Yucatan. It reaches across the Isthmus of Tehuantepec and may
be sympatric with the related species, L. salvini. It is probably the
most common Laccophilus in the southeastern United States. Its dis-
tribution in the northern tier of the United States is in scattered, widely
separated localities that probably represent true distribution and not
collecting gaps. As yet, it has not been found in northern Veracruz;
but additional collecting should produce enough localities to show
that the southern Mexican population is continuous with those in
northern Mexico and Texas (Fig. 7).

Young (1954) describes proximus as one of the principal pioneer
species of newly formed ponds, puddles, and other bodies of fresh
water. He has taken adults in rain barrels, tin cans partly filled with
water, flooded furrows in recently plowed fields, and even in water in
old automobile tires in vacant lots. Throughout the Florida penin-
sula, it is the most common Laccophilus of temporary pools and
puddles. It may appear in large numbers in recently filled, formerly
dry basins. Zimmerman (1960) in Indiana found that proximus
appeared but sporadically in four continuously observed ponds and
did not occur during several months of the year. It was the first of
four Laccophilus species to reappear in ponds that had previously
dried up.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ALABAMA. Mobile County. Chicka-
saw, 1, vi.25.31, H. Dietrich (CNL). Mobile, 1, viii.10.—, Shoemaker
(USNM). ARKANSAS. Garland County. Hot Springs, 5 m. S., 3 ¢, 2 2,
x.17.56, JRZ (NMSU). Hempstead County. Hope, 6, iii.20-27.22, E. W.
Mank (CNL); 1 ¢, vi.20.32, L. Knoble (CAS); 1 m. N., 1 6, 1 2, x.17.56,
JRZ(NMSU). Hot Springs County. County, 3 ¢,3 2, x.17.56, JRZ (NMSU).
Monroe County. Brinkley, 4 6, 6 2, vi.24.48, C. & P. Vaurie (AMNH).
Pulaski County. Little Rock, 2 2, Wickham (AMNH). COLORADO. Sedg-
wick County. Ovid, 4 2, 6 9, viii.17.61, JRZ (NMSU). CONNECTICUT.
Fairfield County. 3 6, vii.4—, C. H. Roberts (AMNH). DISTRICT OF
COLUMBIA. Washington. Shaw Pond, 1, iv.14.26, H. S. Barber (USNM).
Washington, 1, vii.21.—, J. D. Sherman, Jr. (USNM). FLORIDA. Alachua
County. Alachua County, 5, 1.6.23, Chamberlain (CNL); 2 2, ix.26.—-; 1,
Ken ASS exe Sal 44 le Pe i6e23) CAIMINE)) = Hogtown; 35 x.24:37, ESN:
Young (CNL); Hogtown Ck., 1 4, 1 2, x.24.37, F. N. Young (AMNH).
Bay County. Springfield, 1 2, x.16.41, F. H. Young (AMNH). Brevard
County. Titusville, 2 ¢, 1 2, xi.8.11 (AMNH). Broward County. Davis, 5,

MEM. AMER. ENT. SOC., 26

96 THE GENUS LACCOPHILUS

vii.27.39; 2 6, 4 9, vil.27.39, F. H. Young (AMNH). Charlotte County.
Punta Gorda, 5 6, 5 9, xi.16.11 (AMNH). Collier County. Naples, 2 6,
4 9, i1i.30.47, J. W. Green (CAS). Dade County. Coral Gables, 1 ¢, iv.6.46,
F. H. Chermock (CAS). Buck Key, 1, Brainard (USNM). Homestead, 5,
iv.8.51, H. & A. Howden (USNM); 2 6, 3 9, vi.11.51; 4 2, v.11.51, O. Bryant
(CAS). Miami, 1, 1.21.34; 1, x.7.33; 2 9, viii.26.37, F. N. Young (AMNH).
Miami Springs, 1 ¢, viii.2.62, B. Benesh (CAS). De Soto County. Arcadia,
Bunker Branch, 4, 11.20.38; 13, iv.6.39, J. C. Bradley (CNL). Duval County.
Jacksonville, 2, iv.21.—, G. M. Greene (USNM); 13, viii—.02 (USNM);
69 (USNM); 2 6, 2 2, Van Dyke (CAS); 19 8, 30 2, vii—.—; 3 6, 1 2,
vil.—.02; 1 6, vii.4.—; 35 6, 39 9, viii—m—, W. S. Genung (AMNH); 2 2,
v.10.—; v.1.—, P. Laurent (ANSP). Franklin County. Dog Island, 1 92,
iv.16.47, F. N. Young (AMNH). Gulf County. Deadpond, Rockbluff, 1,
iv.3.27, M. D. Leonard (CNL). Wewahitchka, 8, iv.6.27, M. D. Leonard
(CNL). Hendry County. Clewiston, 10, iv.6.44, J. G. Needham (CNL).
Hernando County. Brooksville, 1 ¢, 3 2, i.20-30.40, Van Dyke (CAS). High-
lands County. Lake Placid, 8, iv.7.45; 2, ii.11.49, J. G. Needham (CNL).
Near Avon Park, 1 9, viii.22.61, T. Morris (SCH). Hillsborough County.
Tampa, 4, iv.21-24.—, Hubbard & Schwarz (USNM); MacDill Field, 1 6, 1 9,
vi.5-12.43, B. Malkin (FM). Lee County. Estero, 1, ii.10.44, J. G. Needham
(CNL). Ft. Myers, 1 6, iii.30.12 (AMNH). Leon County. 1, vii.26.22,
Chamberlain (CNL). Martin County. Stuart, 1 6, 2 2, vi.25.51, O. Bryant
(CAS). Okeechobee County. Okeechobee, 5, iii.18.43, W. Proctor (CNL).
Sebring, 1 6, 2 92, vii.20.42, C. Parsons (CAS). Orange County. Winter
Park, 2, ii.7.28; 16, 1.12.29, Gehring (CNL). Palm Beach County. Jupiter,
4, 11.23.43, W. Procter (CNL); 2 ¢, vili.11.43, W. Procter (AMNH). Lake
Worth, 1 6, 4 2 (AMNH). Palm Beach, 3, iii.25.43, W. Procter (CNL);
14,1 9, iv.13.23, R. Hopping (CAS). Pinellas County. Belleair, 1 ¢ (AMNH).
Dunedin, 1, iv.3.23; 3, iv.2-4.23, E. W. Mank (CNL). Polk County. Lake-
land, 3 6, xi.10.11 (AMNH). Putnam County. Crescent City, 6 (USNM).
Palatka, 1, v.3-4.16, J. C. Bradley (CNL). St. Johns County. Hastings, 38,
v.—.—; 38 (USNM); 1, viii.16.50, P. J. Spangler (USNM); 1 (CNL); 2 ¢,
2 2 (AMNH). St. Augustine, 1, C. W. Johnson (USNM). Sarasota County.
Englewood, 1, xii.2.43; 1, iti—.44, J. G. Needham (CNL). Sarasota, 13,
viii.14.10, J. C. Bradley (CNL); 1, ii.7.45, J. G. Needham (CNL); 1 4, ix.14.10,
Van Dyke (CAS). Seminole County. Sanford, 2; 2, vii—m — (USNM);1 4,
2 9,iv—— (AMNH). Taylor County. County, 6 ¢, 12 2, viii— —, W. S.
Genung (AMNH). Volusia County. Enterprise, 1, v.24.—, Hubbard &
Schwarz (USNM); 5 4, 3 9, v.17.—, J. W. Green (CAS). Ormond, 1 3
(AMNH). GEORGIA. Baker County. Elmodel, 1 6, 1 2, x.29.38, H. H.
Hobbs & F. N. Young (AMNH). Bullock County. Statesboro, 1 2, viii.15.50,
F. N. Young (AMNH). Charlton County. Okefenokee Swamp, Billy’s Island,
2, vi—.12, J. D. Sherman, Jr. (USNM); 23, vi—12; 2, v.9.11 (CNL); 1 6,
1 2, Van Dyke (CAS); Mixon’s Hammock, 2, vi.16.12; Tralers Hill, 1, vi.22.—
(CNL). Chattahoochee County. Ft. Benning, 1 ¢, vi.11.43, D. E. Beck

JAMES R. ZIMMERMAN 97

(CAS). Decatur County. Bainbridge, 1, vii.15-27.09; 2 6, ix.17.—; x.19.10,
J. C. Bradley (CAS). Spring Creek, 1, vii.16-29.12; 3, viii.26-28.13 (CNL).
Floyd County. Cave Spring, 1, viii.30.09 (CNL). Fulton County. Ft. Mc-
Pherson, 1 6, 1 9, ix.12.43, D. E. Beck (CAS). Hill County. Gainsville, 6,
iv.2.11 (CNL); 1 4, iv.2.11, Van Dyke (CAS). Jefferson County. Wrens,
1, 11.8.11, J. D. Sherman, Jr. (USNM); 2, iii.8.11 (CNL). Liberty County.
Riceboro, 2 6, 1 2, viii.16.50, F. N. Young (AMNH). Lynn County. St.
Simon Island, 2; 1, ii.16.11; vi.12-22.11, J. C. Bradley (CNL). Pierce County.
Blackshear, 6, v.10.11 (CNL). Offerman, 6, iv.22.11 (CNL). Rabun County.
76,3 2, vii——, C. W. Leng (CAS). Thomas County. Thomasville, 7,
xii.22.34; D. Dunavan (USNM). ILLINOIS. Alexander County. Olive
Branch, 2 2, x.8.09, Gerhard (FM). Champaign County. Urbana, 2, vii.29.07;
2, xi.19.—, J. D. Sherman, Jr. (USNM); 1 6, 1 2, xi.19— (AMNH). Cook
County. Edgebrook (Morton Grove), 13, vii.l15.11 (USNM). Pike County.
Pittsfield, 1 6, viii.17.46, B. Cadwell, 1 2, vii.7.46; 1 2, vii.8.46, A. T. McClay
(DAV). INDIANA. Clinton County. Kirklin, 3 m. W., 5 6, 2 2, ix.26.58,
JRZ (NMSU). Huntington County. Huntington, 2 m. N., 1 2, x.26.56, JRZ
(NMSU). La Porte County. La Porte, 3 2 (CNG). Madison County.
Anderson, 2 6, 2 2, vii.6.57, JRZ (NMSU). Marion County. Indianapolis,
1 3, viil.9.60, JRZ (NMSU). Monroe County. Bloomington, 2, xii.31.49,
F. N. Young (CNL); 2 ¢, 2 9, xii.31.49, F. N. Young (AMNH); 4-5 m. N.,
4 Oi, WellSa08 3 Oy © 85 bc AN Sos © @, 4 O, costes IG, me OR AS, 3 OF
iv.20.56; 2 6, 4 9, iv.26.56; 1 6, v.11.56; 1 2, v.12.56; 2 6, 2 2, vii.21.56;
Onde l4s oe Wiis O0 253 15) 65 7 2 ixcli7-sOy 1556, 6 2, 1x21-56; 7 6, 12 9,
1x.28.56; 2 6, x.30.56; 1 6, x.31.56; 1 6, ii.22.57; 1 9, vi.18.57, JRZ (NMSU).
Owen County. Richland Ck., near Freeman, 1 6, xi.2.55, JRZ (NMSU).
Porter County. Dune Park (Ind. Dunes St. Pk.?), 1, J. D. Sherman, Jr.
(USNM). Posey County. Mt. Vernon, 5 m. E., 3 4, ix.9.56, JRZ (NMSU).
IOWA. Hamilton County. Little Wall L., 1 2, vi.14.52, P. J. Spangler
(USNM). Lee County. Ft. Madison, 2, J. B. Smith (USNM). Osceola
County. Sibley, 1, Shimek (USNM). Polk County. Herrold, 1, vii.24.19,
E. D. Quisfeld (CNL). Storey County. Ames, 1 ¢, v.22.31, G. Hopping
(CAS). Warren County. Indianola, 1 2, vi.14.52, P. J. Spangler (USNM).
KANSAS. Atchison County. Atchison, 1 6, ix.23.56, J. W. McReynolds
(CAS). Cherokee County. Galena, 5, viii.26.04; 15, J. D. Sherman, Jr.
(USNM); 4, viii.26.04 (CNL); 15 (CNL). Dickenson County. Hope, 4 m.
S., 4 6, 2 2, vil.8.60, JRZ (NMSU). Douglas County. Baldwin (City?),
2 6,1 2,J.C. Bridwell (CAS). Lawrence, 12 6,15 2, x.2.51, P. J. Spangler,
Jr. (USNM); 1 6 (MCZ). Lone Star Lake, 2 ¢, 1 2, ix.30.51 (USNM).
County, 1, iv.15.52, P. J. Spangler, Jr. (USNM); 2, F. H. Snow (CNL); 1 ¢,
3 2, x.7.48, J. G. Rozen (BERK). Harper County. Danville, 15 m. NE.,
1 6,3 2, vii.20.60, JRZ (NMSU). Harvey County. Newton, 2 m. SE., 5 ¢,
4 9, vii.21.58; 2 m. E., 4 4, 3 @, vii.6.60; JRZ (NMSU). Johnson County.
Gardner Lake, 2 ¢, 3 2, v.3.52, Bell & Spangler; Sunflower, 1 2, ix.17.52,
P. J. Spangler, Jr. (USNM). Kingman County. Waterloo, 5 m. E., 3 2,

MEM. AMER. ENT. SOC., 26

98 THE GENUS LACCOPHILUS

vii.14.60, JRZ (NMSU). Leavenworth County. Tonganoxie, 1 2, vi.1.52;
3 6,5 9, ix.4.52, P. J. Spangler, Jr. (USNM). McPherson County. McPher-
son, 2, J. D. Sherman, Jr. (USNM); 1 2, W. Knaus (AMNH). Pottawatomie
County. Onaga, 5, J. D. Sherman, Jr. (USNM); 1 6 (AMNH). Reno
County. Haven, 8 m. E., 4 6, 2 2, vii.13.60, JRZ (NMSU). Medora, 1
(USNM); 1°, W. Knaus (AMNH). County, 4, ix.21.11, G. M. Greene
(USNM). Riley County. 5, iv.9.—, Popenoe (USNM). Sedgwick County.
Kechi, 5 6, 12 9, viii.10.59; near Viola, 3 4, 3 2, viii.10.60; Sunnydale, 5 m.
E., 4 6, 1 Q, vi.13.58; Wichita, 3 m. E., 41 6, 46 2, ix.28.57, IRZ (NMSU).
Seward County. Liberal, 2 6, 4 @, viii.12.58; 3 m. NE. 1 6, 2 9, viii.12.58,
JRZ (NMSU). Shawnee County. Topeka, 2, v.24.—; 1, vii.19.—; 1, vii.25.—;
3, Popenoe (USNM). Sumner County. WHunnewell, 5 m. S., 1 3, vi.22.60;
South Haven, 4 m. N., 1 2, vi.23.60, JRZ (NMSU). LOUISIANA. Acadia
County. Crowley, 1, ix.30.11, E. S. Tucker (USNM). Rayne, 3, vi.20.17
(CNL). Cameron County. Cameron, 2 2, vii.10-14.05, Van Dyke (CAS).
Lafayette County. Lafayette, 1 ¢, v.19.51, O. Bryant (CAS). Natchitoches
County. Vowell’s Mill, 2, J. D. Sherman, Jr. (USNM); 4 6, 2 9, Leng
(CAS); 4 6, 4 2, Leng (AMNH). Orleans County. New Orleans, Camp
Planche, 3 9, ii.10.44, iii.10.44, D. E. Beck (CAS). Plaquemines County.
Port Sulphur, 2 °, xi.6.43, D. E. Beck (CAS). St. Landry County. Opelousas,
3 6,1 2, v.—.— (CNG); 4 m. W., 42 6, 64 9, vi.26.63, J. G. & K. C. Rozen
(AMNH). Vermilion County. Gueydan, 33, vi.15-16.25; 46, vii.2.25; 1,
vi.25.26, E. Klambach (USNM). Vernon County. Camp Polk, 1 ¢, vii.—.45,
K. L. Maehler (CAS). Baton Rouge County. Baton Rouge, 1 2, vi.3.64, R.
Hepburn (CAS). MASSACHUSETTS. Middlesex County. Lowell, 1 2,
iv.20.71 (MCZ). MARYLAND. Baltimore County. Baltimore, 1 6, 1 2,
vii.1.—:; 1 2, vii.13.—, F. E. Blaisdell; 1 3, 1 9, vii.1.—; 1 92, vii.13.09, F. E.
Blaisdell (CAS). Montgomery County. Piney Point, 4, viii.26.46, R. I. Sailer
(USNM). MICHIGAN. Berrien County. Warren Dunes St. Pk., 1 ¢, 1 2,
vili.19.52, P. J. Spangler, Jr. (USNM). Emmet County. Pellston, 4 m. N.,
1 Q, vi.25.52; 1 4, viii.6.52, P. J. Spangler, Jr. (USNM). Macomb County.
Mt. Clemens, 1 ¢, vi.24.44, B. Malkin (FM). Marquette County. Marquette,
1, vii.2.16, J. D. Sherman, Jr. (USNM). Van Buren County. South Haven,
12, viii.2.—: 7, viii.11.04, J. D. Sherman, Jr. (USNM); 4 ¢, 5 9, viii.2.—,
V. E. Shelford (AMNH). MINNESOTA. Hennepin County. Excelsior, 3,
J. D. Sherman, Jr. (USNM). Olmsted County. Rochester, 17, vii.12.—; 16,
J. D. Sherman, Jr. (USNM); 3 6, 1 2, vii.l12.—, C. N. Ainslie (AMNH).
County, 2, J. D. Sherman, Jr. (USNM). Ramsey County. St. Paul, 2,
vii.—.—, J. D. Sherman, Jr. (USNM). MISSISSIPPI. George County. Luce-
dale, 1, vii.2.30: 1, vi.22.31, H. Dietrich (CNL). Harrison County. Biloxi,
10 m.N., 2 4,3 9, vili.30.60, P. M. Marsh (DAV). Jackson County. Horn
Island, 1 ¢, v.1.44; 1 4, vi.13.44; 1 6, 1 9, vi.16.44, E.A.R. (USNM). Perry
County. Beaumont, 7, iv.19.32, H. Dietrich (CNL). MISSOURI. Boone
County. Ashland, 4 6, 6 9, ix.9.54, P. J. Spangler (USNM). Columbia, 40
(USNM): 1, iii.29.36, W. M. Gordon (CNL): 1 6, x.8.54, P. J. Spangler

JAMES R. ZIMMERMAN 99

(USNM); 1 @, viii.12.46; 1 2, xii.8.46, W. R. Enns (CAS). Rocheport, 1 m.
W., 16, 1 9, ix.25.54 (USNM). Gentry County. Albany, 2 6, 4 9, vi.13.52,
P. J. Spangler (USNM); 3 6, 4 9, vi.13.52, P. J. Spangler (CAS). Mississippi
County. Charleston, 1 m. N., 1, iv.23.56, P. J. Spangler (USNM). Ripley
County. Doniphan, Logan Ck., 1, iv.23.56, P. J. Spangler (USNM). _ St.
Louis County. Howard Bend, 4, vi.28.37; 17, vii.17.37, W. M. Gordon (CNL).
St. Louis, 2 $, 2 2; 1 4, vi—— (CNG); 4, vii—.77 (USNM). County, 3,
ix.18.40, W. M. Gordon (CNL). Shannon County. Winona, 14 m. S., 1,
iv.—.56, P. J. Spangler (USNM). Vernon County. Nevada, 1 6, v.11.58,
J. W. McReynolds (CAS). NEBRASKA. Arthur County. Near Tyron, 7 é,
3 2, vili.17.61, JRZ (NMSU). Keith County. North of Ogalla, 1 4, 2 2,
vili.17.61, JRZ (NMSU). Lancaster County. Lincoln, 1, Shimek (USNM).
Lincoln County. North Platte, 1 m. S.,2 6, 2 9, vili.16.61; 1 m. E.,3 6,5 @,
Viii.17.61; 10 m. W., 3 6, 5 9, viii.17.61, JRZ (NMSU). Otoe County. Pal-
myra, 1 2, W. F. Rapp, Jr. (WFR). Saunders County. Ceresco, 1, viii.25.59,
W. F. Rapp, Jr. (CAS). NEW HAMPSHIRE. S(outhern?), 1 6, 1 2, x.16.06
(CAS). NEW JERSEY. Bergen County. Fort Lee, 1, ix.22.—, J. D. Sher-
man, Jr. (USNM). Camden County. Berlin, 1 2, vi.24.33, L. J. Bottimer
(BERK). May County. Anglesea, 1 2? (AMNH); 3, G. M. Greene (USNM);
3, C. Boerner (CNL); 1 6, 2 2, v.10.—, J. W. Green (CAS); 4 6, 3 2,
v.8.— (ANSP). Ocean County. Lakehurst, 6, ix.2.—; 1, ix.1.01; 8, J. D.
Sherman, Jr. (USNM); 2 6, 1 2, iv.20.— (CAS). NEW MEXICO. Colfax
County. Cimarron, 10 m. N., 1 2, vii.15.65, A. H. Smith; near Maxwell, 2 2,
viii.14.61, JRZ (NMSU). Harding County. Mosquero, 10 m. E., 2 4, 4 2,
x.23.65, A. H. Smith (NMSU). San Miguel County. Mosquero, 19 m. W.,
1 3, 1 2, x.23.65; 2 m. W., 1 6, 1 2, x.23.65; Trujillo, 19 m. E., Trementina
Ck., 2 2, x.23.65, A. H. Smith (NMSU). NEW YORK. Nassau County.
Flushing, 2 4, viii.11.46, P. Vaurie (AMNH). Niagara County. Olcott, 3,
vili.14.27, H. Dietrich (CNL). Richmond County. Staten Island, 2 (USNM);
3, J. D. Sherman, Jr. (USNM); 1 6, ix.27.38 (FM);5 6, 1 9, ix.11.—, C. H.
Roberts (AMNH). Westchester County. Peekskill, 2; 1, v.12.07, J. D. Sher-
man, Jr. (USNM). Long Island, 2 (USNM). NORTH CAROLINA. Co-
lumbus County, 1, vii.14-21.52 (USNM). Pender County. Willard, 1, v.11.53,
W. M. Kulosh (USNM). Swain County. Bryson City, 8 m. SW., 3 2,
v.18.57, JRZ (NMSU). OHIO. Athens County. Athens, 4 ¢,5 2, v.21-22.51,
P. J. Spangler (USNM). Vinton County. Lake Hope, 1 6, 1 2, vii.10.58,
W. C. Stehr (CAS). Warren County. Mason, 2 m. S., 1 2, iv.24.57, JRZ
(NMSU). OKLAHOMA. Kay County. Braman, 2.5 m. NW., 7 ¢, 7 2,
vi.28.60; Tonkawa, 1 m. S., 2 6, 1 92, vi.23.60, JIRZ (NMSU). Noble County.
Perry, 4m. E., 1 6, 3 Q, viii.1.60, JRZ (NMSU). Pittsburg County. Savan-
nah, 5 4, 5 9, x.19.56, JRZ (NMSU). Seminole County. Seminole, 5 m.
NE., 1 6,2 2, x.19.56, JRZ (NMSU). PENNSYLVANIA. Allegheny County.
Aspinwall, 1 6, ix.18.28; Pittsburgh, 2 ¢, v.10.—; 1 6, 1 2, v.17.25 (CNG).
Butler County. Evans City, Ash Stop, 1 ¢, vii.21.27 (CNG). Dauphin County.
Linglestown, 1, Kaeber (USNM). Elk County. Caledonia, 4, J. D. Sherman,

MEM. AMER. ENT. SOC., 26

100 THE GENUS LACCOPHILUS

Jr. (USNM). Lancaster County. Lancaster, 1, vii.l4.—, Kaeber (USNM).
Philadelphia County. Philadelphia, Phil. Neck, 2, J. D. Sherman, Jr. (USNM);
1 $, x.9.— (FM); 1 2, v.12.— (ANSP). SOUTH CAROLINA. Beaufort
County. Beaufort, 3 6, 1 Q, iii.23.92; 10 3, 13 9, iii.31.92; 2 3,5 9, iv.7.92;
2 6, iv.11.92; 10 $6, 23 9, iv.12.92; 4 2, iv.20.92; 5 6, 8 2, G. D. Bradford
(AMNH). Colleton County. Ruffin, 1, viii.30.28, D. Dunavan (USNM).
Pickens County. Clemson College, 2, xi.22.28; 1, 12.——, D. Donavan
(USNM). SOUTH DAKOTA. Brule County. Chamberlain, 1 4, 1 2, i1x.9.39,
G. B. Spawn (AMNH). TENNESSEE. Shelby County. Memphis, 1 °,
v.11.55, H. F. Howden (DAV). TEXAS. Bastrop County. Bastrop, 1 °,
vii.12.37 (TAM); St. Pk., 1, vi.19.56, Evans & Matthews (CNL). Bexar
County. Leon Ck., 2 2, x.11.52, B. J. Adelson (CAS). San Antonio, 1 ¢,
1 2, iv.9.58, J. F. Laurence (BERK); 1 2?, vi—.42, E. S. Ross (CAS);
1 2, Wickham (CNG). Blanco County. Cypress Mill, 2, Chttn. (USNM);
1 6, ix.20.88; 3 ¢, 1 2, x.20.88 (CAS). Shovel Mtn., 12 (USNM);1 6, 1 2
(CAS); 4 6, 4 2, ix—.—, F. G. Schaupp (AMNH). Bowie County. Tex-
arkana, 2 2, x.17.56, JRZ (NMSU). Brazos County. College Station, Wyxon
L., 2 2, iv.18.53 (TAM): 5 6, 1 2, x.18.56, JRZ (NMSU); Fish Lake, 34 ¢,
48 2, x.17.64-1.24.65, Conte (TAM). Brewster County. Marathon, 2 ¢,
2 2, x.12.49, O. Bryant (CAS). Burnett County. 7, Hubbard & Schwarz
(USNM); 2, J. D. Sherman, Jr. (USNM). County, 1 é, 12 (AMNH).
Calhoun County. Port La Vaca, 1 6, 1954, P. A. Glick (CAS). Cameron
County. Brownsville, 2, v.3.04, H. S. Barber (USNM); 1 (USNM); 1, vii——,
Wickham (USNM): 1 é (FM); 1, vi—.—, Dretz (USNM): 5 (USNM);
146,12, vii——, F. H. Snow; 2 2, x.—.42; 3 6, 1 2, xi.27.42; 1 6, vii.24.42,
E. S. Ross (CAS); 1 ¢, 1 2, ix.22.24, Weed & Pray (FM); 1 4, iv.26.56,
P. A. Glick; 4 2, x.4.57, W. Miller (SCH); Esperanza Ranch, 5, viii.19.—
(USNM); Old Ft. Brown, 1, viii.3.06, A. B. Wolcott (USNM). Seabrook,
1 é, x.20.49, O. Bryant (CAS). Colorado County. Borden, 1, vi.18.—, Hub-
bard & Schwarz (USNM). Columbus, 5, vi.18.—; viii.1.—; v.4.—; vii.24.—;
vi.27.—, C. V. Riley; 2, viii.1-6—, Hubbard & Schwarz (USNM). Comal
County. San Marcos, 13 m. W., 1 &, vi.24-25.61, R. L. Westcott (LACM).
Dallas County. Dallas, 1, v.22.05, I. C. Crawford; 1, v.7.03, C. R. Jones; 3,
Wickham (USNM); 4 6, 22 (AMNH); 1 2, Wickham (ANSP). Duval
County. San Diego, 1, v.23.—, Hubbard & Schwarz (USNM). El Paso
County. El Paso, 1 é (CAS); 2 6, 22 (AMNH). Falls County. Little
Brazos R. at St. Rd. 6, 11 2, 7 2, x.18.56, JRZ (NMSU). Gonzales County.
Harwood, 1, Wickham (USNM). Grayson County. Lake Texoma, below the
dam, 4 6, 1 2, x.19.56, JIRZ (NMSU). Sherman, 17 ¢, 17 @, vi.14.57, D. R.
Lauck (USNM). Grimes County. Singleton, 2m. N., 2 6, 3 2, x.18.56, JRZ
(NMSU).. Harris County. Houston, 11, vi.15.32, Harwood; 1, vii.1.39, Vick
(CNL). Seabrook, 1 2, viii.6—, J. W. Green (CAS). Harrison County.
Karnack, 6 2, vi.17.58, R. L. Fischer (MCHS). Hays County. Dripping
Springs, 1 ¢, 1 2, vili.9.42, W. S. & E. S. Ross (CAS). Hidalgo County.
Edinburg, 11, ii.23.36, J. G. Needham (CNL). Hopkins County. Sulphur

JAMES R. ZIMMERMAN 101

Springs, 1 6, 2 2, x.17.56, JRZ (NMSU). Davis County. Davis Mtns.,
Elbow Can. Crk., 1 6, 3 2, x.27.61; Madero Can., 1 2, x.27.61; Davis Mtns.,
4 46, x.27.61, JIRZ (NMSU). Jefferson County. Pt. Arthur, 3 4, 4 2, x.18.48,
O. Bryant (CAS). Jim Wells County. Alice, 8 m. NE., 9, viii.25.35, C. E.
Burt (USNM). Kerr County. Kerrville, 10, i.22.07, F. C. Pratt (USNM).
Kleberg County. Kingsville, 34, C. T. Reed (CNL); 3 6, 4 2, x.20.61, B.
McDaniels (NMSU). Riviera, 1, ii.28.36,J.G. Needham (CNL). Lee County.
Fedor, 3 6, 2 2, v—.— (CNG). Lexington, 1, x.—.09, Berkiman (CAS).
County, 1, J. D. Sherman, Jr. (USNM); 1, v.—.10, Shoemaker (USNM);
1 6,3 2, ix—.09 (CAS). McLennan County. Waco, 12 m. S., 2 2, x.18.56,
JRZ (NMSU). Nueces County. Corpus Christi, 1, v.10.44, F. R. DuChanois
(USNM). Randall County. Palo Duro St. Pk., 1 4, viii.7.61, H. R. Burke
(TAM). Refugio County. Refugio, 1 2, vi.9.60, F. N. Young (UMMZ).
Sutton County. Sonora, Dry Devils’ River, 1 6, xi.5.49, O. Bryant (CAS).
Titus County. Mt. Pleasant, 5 m. S., 2 3, 4 2, x.17.56, JRZ (NMSU). Travis
County. Austin, 2, vi.28.—, Wickham (USNM); 3 4, 3 2, xi.16.28; 3 6, 2 2,
xli.5.28; 1 6, xii.10.28; 1 6, iv.15.24, J. O. Martin (CAS). Val Verde County.
Del Rio, 2 2, x.5.49, 1 6, xi.5.49, O. Bryant (CAS). Victoria County. Lo-
lita, 1, vii.6.16, J. D. Mitchell (USNM). Victoria, 1, iii.20.01; 2, vi.10.14;
6, ix.18.14; 1, viii.25.15, J. D. Mitchell (USNM) Washington County. Bren-
ham, 1, iv.16.07, R. A. Cushman (USNM). Zapata County. Zapata, 5, 11.26.36,
J. G. Needham (CNL). VERMONT. Bennington County. County, 2 6, 2 2,
vili.16.—, C. H. Roberts (AMNH); 1 ¢ (ANSP). VIRGINIA. Bath County.
Warm Springs, 1 6, x.1— (CAS). Fairfax County. Falls Church, 1, viii.—.14,
St. George & Craighead (USNM). Hansemond County. Suffolk, 2 2, vi.27.76,
Dimwock (MCZ). Stafford County. Fredericksburg, 1, vili.19.89; 2, iii.15.91;
2, 11.26.99; 1, ix.21.00, W. D. Richardson (USNM). WEST VIRGINIA.
Greenbriar County. White Sulphur Springs, 1, vii.23.10, W. Robinson (USNM).
WISCONSIN. Dane County. Madison, 1 4, ix.31.35, H. R. Doge (CAS).
Dodge County. Beaver Dam, 1, vi.22.10, W. E. Snyder (USNM). Sauk
County. Sauk City, 1, J.D. Sherman, Jr. (USNM). Shawano County. Clover-
leaf, 1, viii.17.39, H. R. Dodge (CAS). WYOMING. Goshen County. Ft.
Laramie, 3.5 m. W., 1 @, viii.19.65; 5.3 m. E., 1 9, viii.18.65; Jay Em, 2.5 m.
N., 1 6, viii.20.65; Lingle, 3.8 m. E., 1 6, 1 9, viii.19.65, H. B. Leech (CAS).

MEXICO. — CAMPECHE. Champoton, 11 m. SW., 7 6, 5 2, xi.27.63;
Chencholli, 1 6, 2 9, xi.28.63; Cuidad de Carmen, 4 m. E., 25 6, 27 2,
xi.27.63, JRZ; Puerto Real, 2 m. W., 13 6, 9 2, xi.25.63, K. L. McWilliams
(NMSU). CHIAPAS. Azufre (near Teapa), 5 6, 3 2, xi.26.63; Ocozucuatla,
GuMeWse las txee63) Meapay 9) ms Ne 12 62, 1) 95 xi-261635 IRZ CNMSU):
COAHUILA. Ramos Arizpe, 2 2, vii.7.63, JRZ (NMSU). OAXACA. La
Ventosa, 50 m. N., 1 6, xii.14.55, J. C. Schaffner (UMMZ). Rd. 185, 15 m.
N. of Junct. of Rd. 190 (about 20 m. N. of Juchitan), 1 ¢, ix.2.63, JRZ
(NMSU). SAN LUIS POTOSI. Agua Zarca (near Platinito), 1 ¢; Antiguo
Morelos, 1 4, iii.23.63; Comoca, Rio Axtla, 1 ¢, iii.23.63; Mante, 1 9, iii.23.63,

MEM. AMER. ENT. SOC., 26

102 THE GENUS LACCOPHILUS

JRZ (NMSU). TABASCO. Villahermosa, 5 m. S., 11 6, 6 9, xi.26.63;
2m.E.,6¢6,102,IJIRZ (NMSU). TAMAULIPAS. Magiscatzin, Rio Guaya-
lejo, 2 ¢, 11.26.63, JRZ (NMSU). San Fernando, Rio Linares, 1 ¢ (det. as
compared with type of confusus by FNY), vi.10.60, F. N. Young (UMMZ).
San Jose, 5, iv—.10, J. D. Sherman, Jr. (USNM). VERACRUZ. Catemaco,
1 6, 1 9, vill.26.62; Coatzacoalcos, 15 m. E., 10 6, 1 2, vili.26.62, JRZ
(NMSU). Cotaxtla, 1 ¢, vi.13.58, D. Candia Z. (CAS). YUCATAN. Pro-
greso, 5m. S., 5 6, 7 Q, xi.24.63; Uxmal, 5 6, 6 9, xi.25.63, JRZ (NMSU).

VIRGIN ISLANDS. — Anagarda, 16 6, 27 9, iii.31.25; St. Croix, 9 6, 17 &,
iv.4.25; St. Thomas, 8 ¢, 7 9, ii.28.25 (AMNH).

PUERTO RICO. — Adjuntas, 1 ¢, 1 2, vi.8-13.15; Arecibo, 1 6, ii.—.99,
Aug. Busck; 1 ¢, ii.11.55, D. L. Lauck; 7 4, 1 2, vii.30-viii.1.14 (AMNH).
Bayamon, 3 6,2 2, i.—.99, Aug. Busck (USNM). Caguas, 3 é, 1 2, v.28-29.15;
Coama Springs, 3 ¢, 4 9, vii.17-18.14; 4 6, 4 9, ii.11.— (AMNH). Fajardo,
44,3 2, ii—.99, Aug. Busck (USNM). Isabella, 7 6, 3 2, 1.4.15 (AMNH).
Manati, 1 9, ii.12.55, P. J. Spangler (USNM). Porto Rico, 1, J. D. Sherman,
Jr. (USNM);1 4 (AMNH). San Sabastion, 1 ¢, 1.25.99, Aug. Busck (USNM).

BRITISH WEST INDIES. — BAHAMA ISLANDS. Abasco Cays, Allans Cay,
2 2, v.9.53, E. B. Hayden; Andros Island, Coakley Town, 1 ¢, 1 2, iv.22.53,
E. B. Hayden; Hatchet Bay, 1 9, iv.2.53, E. B. Hayden & L. Giovannoli; Cat
Island, 1 é, 1 @, iii.21.53, E. B. Hayden; Crooked Island, Landrail Pt., 2 ¢,
2 2, 1i.5.53, E. B. Hayden; Eleuthera Island, Governors Harb., 1 ¢, iii.31.53.
Exuma Cay, 1 2, 1.14.53, E. B. Hayden & L. Giovannoli; Grand Bahama, West
Ind., 4 6,2 9, E. B. Hayden; Grand Turk Island, 9 9, ii.19.53, E. B. Hayden;
Great Inagua Island, Matthew, 12 m. N., 1 9, vi.29.53, E. B. Hayden, L.
Giovannoli; Long Island, Deadman’s Cayon, 1 ¢, 1 9, iii.11.53, E. B. Hayden
& L. Giovannoli; N. Bimini Island, 1 ¢, vii—.51, M. Cazier, C. & P. Vaurie;
New Providence Island, Nassau, 1 ¢, 1.3.53; 1 9, iv.5.53, E. B. Hayden
(AMNH); 3 6,42 (ANSP). Rum Cayon, 2 6, 4 2, ili.16.53, E. B. Hayden;
S. Bimini Island, 3 ¢, 4 9, vii— 51, M. Cazier, C. & P. Vaurie; 1 2, v.—.51,
Cazier & Gertsch; 1 2, vili.10.51; C. & P. Vaurie; South Caicos Island, 1 ¢,
2 2, 11.11.53, E. B. Hayden & G. B. Rabb (AMNH).

CUBA. — Havana, 3 6, 1°, Baker (CNG). Jutinica, Alto Sonto, 2 2,
xi.9.24; Pinar del Rio, 1 ¢, 3 @, ix.9-24.13; 10 km. S. Pinar Rio, 2 é, 3 Q,
ix.12-22.13; 7-14 km. N. Vinales, 6 ¢, 13 9, ix.16-22.13; Cabanas, 5 9, 2 4,

ix.5-8.13 (AMNH).
Laccophilus salvini Sharp (Figs. 62-69, 291-292)

Laccophilus salvini Sharp, 1882a, p. 291. Holotype: male, British Museum
(Natural History), Guatemala, Guatemala; Sharp, 1882b, p. 10; Zimmer-
mann, 1920, p. 25; Blackwelder, 1944, p. 74.

DIAGNOSIS. — The combination of posterior elytral blotch, black venter,
sawlike ovipositor, and file in the male and female should separate salvini from

JAMES R. ZIMMERMAN 103

6 doy % , “oa
Ve a
+ LACCOPHILUS PROXIMUS ‘“%

LACCOPHILUS SALVINI 2S:

Figure 7. Distribution of Laccophilus proximus and L. salvini.

MEM. AMER. ENT. SOC., 26

104 THE GENUS LACCOPHILUS

salvini

30 salvini

below if

Tehuantepec yp

3.8 4.0 4.2 4.4 46 48
WOTAE” SEIN GasA nm)

Figure 8. Histograms of length in Laccophilus proximus and L. salvini.
Males are shown crosshatched; females, stippled.

JAMES R. ZIMMERMAN 105

all other North American Laccophilus. It differs from mexicanus and pseudo-
mexicanus in its smaller size (average length, 4.15 to 4.30 mm as compared to
average lengths of over 4.7 mm) and black elytral blotch, and from fuscipennis,
fasciatus, and proximus by the black venter. Small females of mexicanus and
large ones of salvini cannot always be reliably separated, since the blotch is
sometimes weak and there is some size overlap. A mid-lateral elytral macula
is almost always well-defined in salvini, but weak in mexicanus. Small speci-
mens of f. fasciatus can be very similar dorsally, but the differently colored
venter permits reliable separation. The aedeagi of proximus and salvini are
similar enough to pose the question of whether they are specifically different.

DESCRIPTION. — Medium (length 3.9 to 4.8 mm; width 2.3 to 2.7 mm),
dark brown, irrorated species; black beneath; metacoxal file prominent in males
and weakly suggested in females; prosternal process short; ovipositor sawlike.
COLOR. Head: pale brownish-yellow above and beneath, slightly darker at
base of pronotum between the eyes; appendages yellow except for reddish-
brown mandibles. Pronotum: pale brownish-yellow. Elytra: pale brownish-
yellow background with irrorated pattern distinctly outlined (figure 291); strong
tendency to completely suffuse and coalesce in the posterior half to form a
nearly complete transverse fascia or blotch in some specimens; apex darkened,
but usually a large clear area immediately anterior to apex; epipleura pale
anteriorly and dark brown posteriorly. Tergite VIII: dark brown to black.
Venter: prosternum, its process, prolegs, and mesolegs pale brownish-yellow;
hind legs and postcoxal processes reddish-brown, darker at edge and on tarsi;
mesothorax, metathorax, and metacoxal plates black; abdominal sternites from
yellowish-brown to reddish-black with all degrees of intermediacy; males usu-
ally darker than females. Genitalia: generally reddish-brown with varying
degrees of yellow. ANATOMY. Méicroreticulation: weakly double on head,
pronotum, and elytra; secondary mesh discernible, but individual cellules still
apparent. Head: supraclypeal seam closely parallel to margin. Pronotum:
WH/PW, 0.70; LP/PW, 0.40. Elytra: epipleural flange small and seldom
present; truncation slight. Venter: coxal file finely present in males, composed
of about 28 to 30 lines; weakly present in females, but too fine to count; lobes
of postcoxal process rounded laterally and projecting well beyond the midline;
male and female last visible abdominal segments rounded, not truncated, and
similar in outline; that of males slightly produced; female with faint groove on
either side forming a marginate edge; crest fairly well-defined; asymmetrical in
males; broad ridge in females; several rugae near the posterior margin; setiger-
ous punctures thickest near the apex. Legs: male pro- and mesotarsi expanded
in a dorsoventral plane; palettes easily visible at 20 power magnification; fifth
tarsal segment of both pair of legs about one and one-half times as long as corre-
sponding fourth; profemoral setae (5 to 6) shorter and finer than mesofemoral
ones (5 to 6). Genitalia: oval plate with long acuminate tip and a well-defined
median crest which has little anterior curvature; weak raised lines on either
side: aedeagus evenly curved, narrowing toward tip; without distinctive knobs

MEM. AMER. ENT. SOC., 26

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JAMES R. ZIMMERMAN 107

or sculpturing; right paramere with blunt apex; apex of left paramere slightly
asymmetrical; ovipositor with about 13 sawlike teeth.

NOMENCLATURAL NOTES. — Confusion in this taxon has
been limited to it being mistaken for L. confusus in Arizona (Leech
1948a). The specimens were somewhat lighter than usual beneath;
and since salvini and proximus (= confusus) are so similar, it is not
surprising that the Mexican synonym was given to those individuals.

VARIATION. — The most apparent pattern variation is the de-
gree to which the posterior part of the elytra has darkened. Some
have a fascia or blotch nearly as complete as that of L. f. fasciatus,
while others have little enough to be confused with mexicanus. The
darkening is stronger in specimens from south of the Isthmus of
Tehuantepec. Arizona specimens have lighter venters than other
salvini and, except for the elytral pattern, are very close to proximus.

Populations from south of the Isthmus are larger than those to
the north. A comparison of mean pronotal widths for males from
Morelos and Las Casas, Chiapas, which are the closest samples on
either side of the Isthmus, give a highly significant difference (Table
4). The samples from Comitan, Chiapas, and Guatemala agree with
the Las Casas population. One could readily justify describing the
northern populations as a different race, but I have not done so. The
population from Arizona appears even more different than that from
Central Mexico, but only ten specimens were available for compari-
son. The species shows considerable geographic variation and de-
serves additional descriptive analysis (Fig. 8).

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— L. salvini occurs from southern Arizona to Guatemala. At pres-
ent there is a gap in the records from northern Sonora to Nayarit.
Too few collections have been made on the west edge of the Sierra
Madre Occidental to consider this more than a collecting gap, how-
ever. It is common in Nayarit, Jalisco, Michoacan, and Morelos,
and further south in Chiapas and Guatemala. It generally is found
between two and seven thousand feet (Fig. 7).

Like proximus it seems to be a pioneer species in newly formed
situations at higher altitudes. It seldom occurs on the coastal low-
land. Grassy margined ponds are the most common habitat.

MEM. AMER. ENT. SOC., 26

108 THE GENUS LACCOPHILUS

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Pima County. Tucson,
Bear Can., 1 6, iii.18.46; 2 6, 3 9, iii.26.46, J. W. Green (CAS). Catalina
Mtns., 2 6, 2 2, iv.18.—, Hubbard & Schwarz (USNM); 1 9, xi.8.34, K.
Thomas (NMSU).

GUATEMALA. — Carcha (Dept. Alto V. P.), 9 6, 2 2, vi.23.66, Flint &
Ortiz; Chimaltenango, 20 m. NW., 48 6, 40 @, viii.20.65, P. J. Spangler;
Guatemala City, 20 m. S., 10 6, 7 9, vii.7.65, P. J. Spangler (USNM).

MEXICO. — CHIAPAS. Buchil (near), 1 ¢, viii.31.63; Comitan, 92 8, 69 2,
viii.30.63; 13 m. W., 38 6, 40 9, viii.30.63; Ixtapa, 15 6, 6 9, viii.31.63;
Ocozucuatla, 6 m. W., 1 6, ix.1.63, JRZ (NMSU). San Cristobal de las
Casas, 1 6, 1 9, 1.25.66, P. H. Raven & D. E. Breedlove (CAS); 38 6, 17 2,
vili.28.63; 5 m. E., 17 6, 7 2, viii.28.63; 20 m. W., 3 6, 1 9, viii.30.63;
Tuxtla Gutierrez, 10 m. W., 6 6, 5 2, ix.1.63, JRZ (NMSU). COLIMA.
Trapechi (near Colima), 2 6, 6 2, vii.29.62; 15 6, 22 9, vii.30.62, JRZ
(NMSU). JALISCO. Ameca, 25 km. E., 43 6, 51 9, iii.25.64, JRZ (NMSU).
Atenquique, 15 m. NE., 3 6, 3 @, xii.5.48, H. B. Leech (CAS). Autlan,
53 6, 38 2, x.26.66, A. H. Smith & JRZ (NMSU). Barra de Navidad, 1 ¢,
ili.18.61, C. O. Morse; Colima, 29 m. NE., 3 6, 1 2, xii.3.48, H. B. Leech
(CAS). Cuidad Guzman, 5m. N.,5 6, 1 Q, vii.28.62; Guadalajara, 20 m. E.,
46,1 9, 11.25.64; 11 m. S., 8 6, 11 9, vii.30.62, JRZ (NMSU). Jiquilpan,
15-20 m. W., 1 6, 1 2, xi.30.48, H. B. Leech (CAS). La Huerta, 6 m. N.,
8 6,8 2, x.25.66, A. H. Smith & JRZ (NMSU). Mazamitla, 17 m. S., 10 ¢,
8 2, xii.5.48, H. B. Leech (CAS); 5 km. E., 1 6, 2 @, iii.27.64, JIRZ (NMSU).
Tala (near), 27 6, 17 2, ii1.25.64; Tamazula, 5 ¢, 6 2, ui.27.64; Tecalitlan,
10 m. S., 6 é, 13 @, iii.27.64, IRZ: Tecolotlan, 12 ¢, 12 2, x.24.66, A. H.
Smith & JRZ; Tehetlican (Rd. 110), 12 6, 12 Q, vii.28.62; Tizapan (near),
Rd. 15, 26 ¢, 17 9, iii.26.64, JRZ (NMSU). Tlaquepaque, 2 2, vii—m.53,
N. L. H. Krauss (CAS). Union de Tula, 10 m. S., 9 &, 18 9, iii.28.64, JRZ
(NMSU). La Venta de Mochitiltic, 5 ¢, 79, vii.20.55, R. B. Selander
(CAS). MEXICO. Near Tonatico, 1 6, 2 2, vili.29.62, JRZ (NMSU).
MICHOACAN. Carapan, 15 m. S., 2 4, xil.7.48, H. B. Leech (CAS). Jacona,
1 9, viii.11.57, D. R. Lauck (USNM). Jiquilpan, 8 km. E., 11 6, 10 2,
1.2.64; 11 m. W., 2 4, vil.28.62; Uruapan, 2 6, 1 2, iti.26.64; 10 m. S.,
10 6, 16 9, i11.26.64, JRZ (NMSU). Zamora, 9 m. W., 3 6, 1 2, xii.6.48,
H. B. Leech (CAS); 2 2, vii.28.62, JRZ (NMSU). MORELOS. Acatipaca,
4 6, 3 2, vili.29.62; Cuautla (near), 10 2, 8 2, viii.28.62, JRZ (NMSU).
Cuernavaca, 14 m. S., 2 4, x1i.8.48, H. B: Leech (CAS); 1 9, iv.15.46, J. D.
Pallister (AMNH); 2 6,2 2 (UMMZ). Puente de Ixtla, 3 6, 2 92, viii.29.62;
Temixco, 3 ¢, 2 9, vili.29.62, JRZ (NMSU). NAYARIT. Ixtlan del Rio,
34 6,21 Q, 1x.22.53; San Blas, 1 6, 2 2, 1x.17-21.53, B. Malkin (CAS); 5 m.
E., 2 6, 1 2, vi.31.62, JRZ (NMSU). Sierra de Zapotan, 1 6, xi.—.42,
Eugenio Paredes (CAS). Tepic, 1 6, ix.15-17.53; 3 é, 3 2, ix.21-24.53;
19 m. SE., 1 6, 7 92, ix.24.53, B. Malkin (CAS); 2 9, xi.27.48; 15 m. SE.,

JAMES R. ZIMMERMAN 109

1 $, 6 2, xi.28.48; 25 m. SE., 6 6, 11 9, xi.23.48, H. B. Leech; 20.5 m. NW.,
10 6,3 9, xi.24.48, H. B. Leech (CAS). OAXACA. Camotlan (near), 3 4,
4 2, ix.3.64; Huajuapan, 2 km. S., 4 6, 2 2, ix.4.64, JRZ (NMSU). PUEBLA.
Amatitlan, 9 m. N., 1 6, xii.10.48, H. B. Leech (CAS). Puebla(?), 1 4,
vi.26.57, D. R. Lauck (USNM). SONORA. Aribaba (near Moctezuma),
1 $6, 1 2, xii.15.62, JRZ (NMSU). Hermosillo, 3 2, vii.9-16.53, B. Malkin
(CAS). Near Rancho Uriquepa, 1 2, vi.26.56, F. N. Young (UMMZ).
ZACATECAS. Fresnillo, 16 m. NW., Rio Trujillo, 6600 feet, 1 2, vi.29.54,
R. H. Brewer (CAS).

LACCOPHILUS MEXICANUS

This polytypic species is composed of three races in North Amer-
ica. It appears to be part of a complex that extends to southern South
America. Laccophilus chilensis Sharp bears strong resemblance to
this group. As yet, no intergrade zones have been found between the
North American races. One suspected intergrade has been seen be-
tween mexicanus and atristernalis. It is a female, however, and does
not permit a reliable determination of intergradation. The races are
very much alike, and individuals of the two races mentioned above
can be separated only on the basis of male genitalia. The southern
race, oaxacensis, does show more differentiation, and details of pat-
tern and size permit good separation between it and mexicanus. The
ranges of the southern races are, apparently, completely allopatric
with a gap of about 200 miles.

DESCRIPTION. — Medium to large (length, 4.2-5.4 mm; width, 2.4-3.1
mm) species, brown above and dark brown or black beneath; metacoxal file
present; prosternal process short; ovipositor sawlike. COLOR. Head: it and
its appendages yellow or light brown, darkening to light reddish-brown near
the pronotum; mandibles also darkening toward tip. Pronotum: yellow to
yellowish- or reddish-brown. Elytra: light yellowish-brown background with
relatively even irrorations, anterior fingerlike detail of maculosus group gener-
ally lacking in mexicanus and atristernalis, but evident in oaxacensis; some
tendency for suffusion and anastomosing of dots in posterior half and near the
medium suture; dots frequently forming chains; suture dark brown; pattern
limits ill-defined; epipleura pale anteriorly, dark in constricted posterior part.
Tergite VIII: dark brown to black. Venter: prosternum, its process, fore and
middle legs light brownish-yellow with reddish tinge; hind legs of about the
same color with much stronger reddish highlights; mesosternum, metasternum,
metacoxal plates, except postcoxal processes, very dark brown or black; ab-
dominal segments variable from brownish-yellow to black; females with abdomi-
nal segments frequently pale except for the black hind margin of first visible
segment, but other segments tend to be suffused with some dark brown or black.

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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AMER. ENT. SOC., 26

MEM.

1e2. THE GENUS LACCOPHILUS

Genitalia: oval plate, aedeagus, and parameres with varying amounts of dark
reddish-brown or yellowish-brown; ovipositor reddish-brown. ANATOMY.
Microreticulation: finely meshed and only weakly double at best (finer and less
impressed than in maculosus) on head, pronotum and elytra. Head: supra-
clypeal seam closely parallel to margin. Pronotum: WH/PW, 0.67 to 0.68;
LP/PW, 0.40 to 0.41. Elytra: epipleural flange rare in atristernalis and mexi-
canus, but common in oaxacensis; truncation of elytral apices slight. Venter:
coxal file prominent in males, composed of from 25 to 30 lines, laterally 10 or
15 more which are much weaker and not distinctly part of the file; a faint trace
of file apparent in females; prosternal process with well-defined crest; lobes of
postcoxal processes rounded and laterally projecting well beyond the midline;
male last visible ventral abdominal segment slightly produced with or without
rugae along hind margin; asymmetrical curving ridge on middle forming de-
pression on left side or both sides; female last segment either subtriangular or
rounded; slight crest in posterior half of segment. Legs: male pro- and meso-
tarsi noticeably enlarged in a dorsoventral plane, easily observed at 20 power
magnification; fifth tarsal segment on pro- and mesolegs from one and three-
quarters to twice as long as fourth; profemoral setae (5 to 6) shorter and finer
than mesofemoral ones (5 to 7). Genitalia: oval plate with acuminate tip and
well defined crest which anteriorly curves to the left; numerous raised lines on
either side of the crest; aedeagus of complex sculpture and abruptly constricting
near the apex; right paramere with small distinct conical apex; ovipositor with
13 to 15 sawlike teeth.

NOMENCLATURAL NOTES. — The biological situation in L.
mexicanus seems to be a relatively uncomplicated problem, but the
nomenclatural condition has been one of continuous confusion. This
is understandable, since the species lacks the pattern and colors that
might lead one to easily recognize and define the geographical races.
Aubé described mexicanus in 1838 from a single specimen from Mex-
ico. No further locality information was given. I have not seen his
type. It is apparently not at the Institute Royal des Sciences Natur-
elles de Belgique in Brussels or in the Paris Museum. It appears to
be lost; but since the writer has not visited the European museums
that might contain the type, a neotype has not been designated in this
paper. I have seen a female specimen from Guanajuato (Brussels
Museum) compared by Regimbart, with the type, and I have also
examined a male in the Paris Museum from the collection of Regim-
bart; from Durango, Mexico; both are what have been considered
mexicanus. Without examination of the type, there cannot be cer-
tainty about the identity of mexicanus. The specimens from Guana-
juato and Durango belong to a race that extends to south of Mexico

JAMES R. ZIMMERMAN 113

City and north into the southwestern United States. Two other races
occur in North America, however.

Crotch (1873) described atristernalis from California. Sharp and
Horn synonomized it with mexicanus. Leech (1948) maintained
that mexicanus specimens from Oaxaca were different from the Cali-
fornia and Lower California specimens. He mentions the elytral
apices as pale in those from Oaxaca and dark in the ones from north
of there. The male aedeagus is also different and supports Leech’s
observations. The specimens from Oaxaca are being described as a
new subspecies. The material from California is heterogeneous, how-
ever, since there are two races in that state. The one in southern and
Lower California is what is here considered to be typical mexicanus.
The one from northern California, Oregon, Idaho, Nevada, and Utah
is being retained as atristernalis. I have not been able to find Crotch’s
type of atristernalis, and California has proven to be too vague a
locality. The type locality is being restricted to Woodside, San Mateo,
California; and the type locality for the nominate race is being re-
stricted to Guanajuato, Guanajuato, Mexico.

Aubé’s mexicanus could have been either of the two races that oc-
cur in Mexico; but since there is no certain way of establishing which
one he had in hand, it seems better to describe as new the race that
has the more restricted distribution and retain mexicanus for the race
that is common over most of the country. There appears to be no
confusion of mexicanus with any other species of North American
Laccophilus.

Laccophilus mexicanus mexicanus Aubé, new status
(Figs. 70-73, 75, 76, 81, 294)

Laccophilus mexicanus Aubé, 1838, p. 420. Holotype: location not known;
type locality here restricted to Guanajuato, Guanajuato, Mexico. Sharp,
1882a, p. 820; Sharp, 1882b, p. 9; Zimmermann, 1917, p. 122; Leng, 1920,
p. 77; Zimmermann, 1920, p. 22; Blackwelder, 1944, p. 74; Leech, 1948b,
p- 401.

DIAGNOSIS. — The almost evenly irrorate elytra, black venter, and size
are the best general characteristics of this race. Only examination of the male
aedeagus will permit certain identification, however. L. spergatus, L. mistecus
mistecus, L. pseudomexicanus, and L. salvini are sympatric with mexicanus and
have the same general coloration and pattern above and beneath. The first
three species are usually larger than mexicanus, and salvini is smaller; but some

MEM. AMER. ENT. SOC., 26

114 THE GENUS LACCOPHILUS

)

LACCOPHILUS M. ATRISTERNALIS

eo

LACCOPHILUS PSEUDOMEXICANUS jest Cy

Qe td we
%

—JI
INS le AL SA

LACCOPHILUS M. OAXACENSIS © —-—\

Figure 9. Distribution of Laccophilus mexicanus and L. pseudomexicanus.

individuals overlap in size. All other species within the range of mexicanus are
brown, reddish-brown, or yellow beneath.

VARIATION. — The most common variation in pattern is the
degree to which the dots tend to fuse. There may be considerable
fusion in the posterior half of the elytra; but unlike salvini or fasciatus,
it is always reddish-brown and not black. The ventral, abdominal
sclerites of females have varying amounts of brown or yellow instead
of black. It is not a constant feature, however.

JAMES R. ZIMMERMAN 115

Size is variable, (Table 5) but there are no geographical trends.
Males of some population samples (Mexico and Chihuahua, for ex-
ample) may exceed an average of over 4.9 mm, while a sample of 25
males from near Guadalajara, Jalisco, average only about 4.64 mm.
Females appear to be slightly smaller than males, with the means smal-
ler for every locality. The peaks of the frequency distributions for
males and females are at about 4.8 mm, but the profile of the males
is displaced toward the right when compared to the females (Fig. 10).

The PW/EL ratio is remarkably uniform in nine localities extend-
ing from Baja California to Michoacan, which supports the general
geographical similarity (Table 5).

GEOGRAPHICAL RANGE AND HABITAT PREFERENCE.
— The northern limits of this race appear to be the southern edges of
Colorado, Utah, and Nevada (Fig. 9). I have not seen any speci-
mens from Nevada, but they should occur there. It also occurs in
southern California from a line about even with Los Angeles County
and Barstow. It reaches the tip of Baja California and probably oc-
curs generally over the peninsula. Leech (1948b) records mexi-
canus in Baja as atristernalis. This race extends no farther south
than about Kern County, California, however. The northeastern
limits of mexicanus are less well known. It reaches southeastern Col-
orado, northeastern New Mexico, and is common in the Davis Moun-
tains and Big Bend region of Texas. There is a dubious record of a
single specimen taken from Edinburg, Texas (J. G. Needham, ii.
24.36, Cornell University Collection), which is about 300 miles out
of the range. In Mexico mexicanus, along with fasciatus, is one of
the most common species on the Plateau. The race reaches Jalisco,
Michoacan, Mexico, and Puebla at about 5000 feet elevation. It also
occurs in Veracruz, interior to the coastal lowlands. It is sympatric
with four other very similar species, all of which may occur within a
few miles of one another, with up to three species in the same pond.
They are salvini, pseudomexicanus, spergatus, and mistecus. Without
males, experience is needed to distinguish among this assemblage.
Apparently, there is some selective factor that has produced the rather
evenly irrorated brown dorsal pattern and the black venter that is
common to these species.

L. m. mexicanus seems to be most common in transitory, grassy

MEM. AMER. ENT. SOC., 26

116 THE GENUS LACCOPHILUS

margined ponds with mud bottoms. It is less common in gravelly
streams. Its most frequent occurrence is above 3500 feet.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Benson,
1 3,1 Q, 11.25.24, O. Bryant (CAS); 2 m. E., 1 2, x.6.63, K. L. McWilliams
(NMSU). Chiricahua Mountain, 1, vi.16— (USNM); SWRS, near Portal,
1 2, vi.18.58, C. D. MacNeill (CAS); SWRS, 2 6, iv.26.56; 2 2, iv.8.56;
1 2, iv.21.56, E. Ordway; 2 6, x.1.55, M. Cazier (AMNH); Herb Martyr
Lake, 1 2, ix.5.61, JRZ (NMSU). Fairbank, San Pedro River, 1, iii.12.13
(CNL). Huachucha City, 5 6, 3 2, ix.6.61, JRZ (NMSU). MHuachucha
Mtns., 3, v.7.53; Bear Canyon, 2, v.8.53, A. & H. Dietrich (CNL). Palomas,
1 (CNL); 2 (USNM). Rodeo, N.M., 2 m. NW., 2 6, ix.5.61; Tombstone,
5 m.N., 2 6, ix.5.61, JRZ; Wilcox, 11 m. W., 1 2, x.6.63, K. L. McWilliams
(NMSU). Coconino County. Grand Canyon, Supai, 3500 feet, 1 2, viii.2.34,
E. L. Bell (AMNH). Graham County. Geronimo, 1 2, iv.28.24, J. O. Martin
(CAS). Maricopa County. Palo Verde, 1, vi.4.19, A. Wetmore (USNM).
Phoenix, 11 ¢, 9 2, Chas. Palm (AMNH). Wickenberg, 1, xii.28.45, H. P.
Chandler (BERK). Pima County. Arivaca, Arivaca Ck., 1 6, 1 9, vii.31.52,
H. B. Leech & J. W. Green (CAS). Quitobaquito, OPCNM, 4, iv.11.53; 4,
v.1.53, A. & H. Dietrich (CNL). Sta. Catal. Mtns., 1 ¢, iv.10.38, F. H.
Parker (DAV). Tucson, 1, ii.4.20 (ARI); 3 6, 2 2, ix—.33; 2 2, vi-——,
O. Bryant (CAS). Santa Cruz County. Nogales, 5 m. NW., 2 9, ix.6.61;
Santa Rita Mtns., Florida Can., 2 6, xii.2.61, JRZ (NMSU). Yavapai County.
Minnehaha, 1 2, Barber (AMNH). Prescott, 4, ——.— (USNM);5 6, 6 2,
viili—.— Barber (AMNH). Yuma County. Fort Yuma, 8, vi.4.— (USNM).
Yuma, 2 6, viii.24.06 (FM). CALIFORNIA. Imperial County Brawley, 3,
viii.8.14; 7, iii—.16, J. C. Bradley (CNL); 2 2, iii—.16 (CAS). Callipatria,
1 4, xi.14.23, R. C. Casselberry (AMNH). Desert Shores, 20 m. S., 2 6,1 &,
iv.24.62, IRZ (NMSU). Salton, 1, viii—.20 (USNM). Westmoreland (near),
1 8, iv.24.62, JRZ (NMSU). County, 1 6, v.24.25 (CAS); 1 6, 1 9, iv.3.24
(CNG). Los Angeles County. Claremont, 2 ¢, 4 2, Baker (CNG). Los
Angeles, 1 6, —.9.19 (CAS). Los Angeles, 3, Coquillet, 1, W. D. Richardson
(USNM). St. Ana Can., 1, ii.27.26, A. J. Muchmore (MCZ). Orange
County. Laguna, 1 9, ix.9.—, F. E. Winters (CAS). Laguna Beach, 1,
—.—.— (CNL). Riverside County. Coachella Valley, 4 ¢, 1 2, v.25.28,
Van Dyke (CAS). Palm Can., Palm Springs, 1 2, vii.13.97, A. P. Morse
(MCZ). Riverside, 3, F. E. Winters (CNL); 2 6, 62, F. E. Winters
(CAS). County, Sonorian (sic!) Region, Colorado River, on some, 3 ¢, 2 2,
F. E. Winters (CAS). San Bernardino County. Barstow, 1 (USNM). Needles,
1 (USNM); 2 6, 2 2, Wickham (AMNH). San Diego County. Escondito,
12 m.N., 1, ii.20.53, R. K. Benjamin (USNM). San Diego, 2 2, viii.19.37,
F. W. Furry (LACM). COLORADO. Huerfano County. Near Walsenberg,
16, 1 9, vili.18.61, JRZ (NMSU). La Plata County. Durango, 14 m. S.,
1 6, vii.28.66, A. H. Smith (NMSU). NEW MEXICO. Bernalillo County.

JAMES R. ZIMMERMAN 117

Albuquerque, 10; 3, vii.2.—; 2, x.18.19 (USNM); 1 6, 12, Wickham (CAS);
646,292,246, H. F. Wickham (AMNH); 4, vii.4— (ISU). Catron County.
Glenwood, 2 6, 2 2, iv.14.65, A. Smith, R. Smith & JRZ (NMSU). Que-
mado, 10 m. N., 3 9, vii.17.63, R. D. Ohmart (NMSU). Reserve, 1 2,
iv.15.65, A. Smith, R. Smith & JRZ (NMSU). Colfax County. Cimarron,
10 m. N., 2 6, vii.15.65, A. H. Smith (NMSU). Maxwell (near), 1 6, 1 2,
vili.14.61, JRZ (NMSU). Dona Ana County. Las Cruces, 1 6, x.1.62, K. L.
McWilliams; 4 ¢, 3 2, vii.6.61, E. E. Staffeldt & JRZ (NMSU). Mesilla,
1 m. SW., 5 6, 7 Q, viii.l1.61 (NMSU); Mesilla, Mesilla Dam, 1 4, iv.25.24,
J. O. Martin (CAS). Mesquite, 1 ¢, vii.10.61, JRZ (NMSU). Grant County.
Cliff, 12 m. S., 5 6, 13 9, iv.14.65, A. Smith & JRZ (NMSU). AHarding
County. Mosquero, 10 m. E., 1 2, x.23.65, A. H. Smith (NMSU). Hidalgo
County. Animas, 1 m. S., 3 6, vii.31.65, H. B. Leech (CAS); 32 m. S., Day
Ranch, 2 4, vii.31.65, H. B. Leech & Van Tassel (CAS); 5 m. S., 1 2, vii.24.61,
JRZ (NMSU); 12 m. SE., 50 6, 70 2, xi.23.62, K. L. McWilliams (NMSU).
Cienega Lake (near Rodeo), 5 9, vii.17.64, R. H. Arnett, Jr. & E. Van Tassel
(CUA). Near Cloverdale (30 m. S. Animas), 5 6, 7 @, vii.24.61; Gary, 5 m.
W., 7 6, 2 2, vii.24.61; Lordsburg, 12 m. W., 7 6, 2 2, vii.24.61; Lordsburg,
12 m. W., 7 6, 6 2, x.6.63, JRZ; 13 m. E., 2 6, 2 2, x.6.63, K. L. McWil-
liams; Roadforks, 2 2, ix.5.61, JRZ (NMSU). Rodeo, 12 m. N., 1 4, ix.5.61,
JRZ (NMSU). Luna County. Columbus, 1 m. S., 9 6, 7 9, vii.17.61, JRZ;
Deming, 16 m. W., 2 6, x.6.63, K. L. McWilliams (NMSU). Mora County.
Mora, 10 m. S., 1 2, v.1.66, A. H. Smith (NMSU). Otero County. Pinon,
40 m. S., 3 4, vii.29.63, R. D. Ohmart (NMSU). Quay County. Tucumcari,
10 m. W., 2 6, ix.10.63, K. L. McWilliams (NMSU). Rio Arriba County.
Abiquiu, 3 6, 1 2, ix.16.49, O. Bryant (CAS); Canjilon, 1, vii.6.30 (AMNH).
Dulce, 1 4, vi.24.65; Ojo Caliente, 1 ¢, vi.23.65, A. H. Smith, R. L. Smith &
JRZ (NMSU). San Juan County. Fruitland, 1 2, xi.14.62; Newcomb, 2 m.
N., 1 6, vil.9.63, R. D. Ohmart (NMSU). TEXAS. Brewster County. Big
Bend Ntl. Park, Boquillas, 1 ¢, viii.2.61; Marathon, 10 m. S., 1 4, viii.1.61,
JRZ (NMSU). County, 1 é, v.3.27, J. O. Martin (CAS). El Paso County.
Clint, 1m: S= 5 65 2.2, vui2.6l, JRZ (NMSU). El Paso; 1, vit.26.14: 1,
vii.28.14, J. C. Bradley (CNL); 3 (NMSU); 3 6, 1 2 (AMNH). Hidalgo
County. Edinburg, 1, 11.24.36, J. C. Needham (CNL). J. Davis County.
Davis Mtns., 1 6, 1 2, x.27.61; Elbow Canyon Ck., 1 6, 1 2, x.27.61, JRZ
(NMSU). Ft. Davis, 6 é, 3 92, iv.19.53, B. Adelson & M. Fashbauer (CAS).
Presidio County. Marfa, 14 m. N., 1 4, vii.19.61, JRZ (NMSU). UTAH.
Washington County. St. George, 1, vi—— (USNM); 3 4, 4 2, vii——,
Wickham (MCZ).

MEXICO. — BAJA CALIFORNIA SUR. Cabo San Lucas, 7.7 m. NE., 1 9°,
1.1.59, H. B. Leech (CAS). El Carrizal, 1 ¢, 3 2, iv.25.47, Ira La Rivers
(BERK). La Paz, 8.2 m. W., 1 6, xii.31.58, H. B. Leech (CAS). Padarone,
Arroyo Amarillo, 2 2, v.11.47; San Bartolo, 1 ¢, 1 2, v.1.47, Ira La Rivers
(BERK). San Jose del Cabo, 1 m. SE., 1 6, 1 2, 1.7.59, H. B. Leech (CAS).

MEM. AMER. ENT. SOC., 26

118 THE GENUS LACCOPHILUS

San Luis Gonzaga, 12 m. E., 2 6, 7 2, v.22.47; Todos Santos, 2 6, 11 9,
iv.10.47, Ira La Rivers (BERK). CHIHUAHUA. Camargo, 1 m. N., 9 é,
15 2, vii.25.62; Casas Grandes, 5 m. E., 9 ¢, 11 2, xii.20.64; Chihuahua,
43 m. N., 31 6, 25 92, xii.8.62; Colonia Juarez, 2 m. SW., 3 6, 2 9, xii.20.64;
Hidalgo d. Parral, 10 m. S., 1 $, 2 9, xii.9.62, JRZ (NMSU). Parrita, 12 m.
W., Santa Clara Can. (in Canon Prieto), 3 2, vi.30.47, M. Cazier (AMNH).
Primavera, 3 2, vi.30.47, M. Cazier, W. Gertsch & R. Schrammel (AMNH).
Sta. Barbara, 1 ¢, v.14.—, G. M. Bradt (AMNH). DISTRITO FFDERAL.
San Jeronimo, 1 é, 1 2, vi.l1.46, J. & D. Pallister (AMNH). Xochimilco,
1 2, iv.26.46, J. & D. Pallister (AMNH). DURANGO. Abasolo, 3 6, 8 2,
vii.25.62, JRZ (NMSU). Durango, 1 6, 29 9, v.30.37, M. Embury (CAS);
16 6, 15 Q, vii.26.62; 10 m. N., 1 6, 1 2, xii.9.62; 20 m. W., 16 6, 12 9,
xii.10.16; Llano Grande, 10 m. E., 1 6, 1 2, xii.10.62; Navios (Rd. 40), 1 ¢,
xii.10.62; Nombre de Dios, 4 ¢, 2 2, vii.26.62; 6 6, 6 Q, vii.13.63; 9 6, 7 Q,
viil.21.63; San Juan del Rio, 3 ¢, 8 2, xii.9.62, JRZ (NMSU). Tapias, 6400
feet, 2 2, vi.20.61, R. A. Schreibner (MCHS). GUANAJUATO. Guana-
juato, 2 6,5 9, vi.20.57, D. R. Lauck (USNM). Irapuato, 1 ¢, 2 2, xii.22.55,
Alan Gillogly (CAS); 5 m. S., 13 ¢, 7 9, vii.27.62; Leon, 5 m. S., 2 6, 4 @,
vii.26.62, JIRZ (NMSU). Near Cuitzeo, 30 m. N. Morelia, 1 ¢, 1 2, D. Breed-
love (CAS). JALISCO. Ameca, 25 km. E., 1 46, 1 9, iii.25.64, JRZ (NMSU).
Atenquique, 15 m. NE., 5 6, 2 9, xii.5.48, H. B. Leech (CAS). Guadalajara,
11m. S., 4 ¢,3 9, vii.30.62, JRZ; Barranca Oblato, 1 2, x.23.66, A. H. Smith
& JRZ; Lagos de Moreno, 1 6, 10 9, vii.26.62, JRZ (NMSU). Mazamitla,
17 m. S., 3 2, xii.5.48, H. B. Leech (CAS); 5 km. E., 4 4, 2 Q, iii.27.64;
Near Tizapan, 11 4, 8 Q, iii.26.64, JRZ (NMSU). Ojuelos de Jalisco, 12 m.
S., 8 6, 14 9, xi.21.48, H. B. Leech (CAS). Tamazula, Junction Rd. 110 &
Rd. to Cd. Guzman, 1 @, i1.27.64, JRZ; Tecolotlan, 6 ¢, 7 2, x.24.66, A. H.
Smith & JRZ (NMSU). 15-20 m. W. Jiquilpan, 11 ¢, 11 2, xi.30.48, E. S.
Ross (CAS). Zapotlanejo, 10 m. E., 9 4, 8 2, iii.25.64, JRZ (NMSU).
MEXICO. Acambay, 14 6, 7 9, ili.27.63, JRZ (NMSU); 8.7 m. NW., 1 ¢,
ii.3.53, I. J. Cantrall (UMMZ). Atlocomulco, 12 ¢, 1 9, iii.27.63; Tenango
de Valle, 1 ¢, 2 2, viii.29.62, JIRZ (NMSU). MICHOACAN. Carapan, 6 é,
12 2, vii.28.62, JIRZ (NMSU); 15 m. S., 3 6, 7 92, xii.7.48, H. B. Leech
(CAS). Cheran, 2 m. N., 2 2, iii.26.64; Ins. Morelos N. Pk., 2 6, 1 2,
vil.27.62, JIRZ (NMSU). Jacona, 5 ¢, 6 9, viii.11.57, D. R. Lauck (USNM).
Jiquilpan, 11 m. W., 1 6, 2 2, vii.28.62; 8 km. E., 14 6, 6 @, ili.27.64,. JRZ
(NMSU). Lago de Patzcuaro, 1 4, iv.—.66, M. Rosas & P. Reyes (CAS).
Morelia, 11 6, 4 92, viii.12.63; 10 m. N., 4 2, 2 2, vii.12.63; Near Cuitzeo,
6 3, 4 9, vii.27.62; Zamora, 8 ¢, 8 Q, vii.28.62, JRZ (NMSU): 9 m. W.,
4 3,7 9, xii.6.48, H. B. Leech (CAS). PUEBLA. Acatzingo, 3 ¢, viii.28.62,
JRZ (NMSU). Lago Alchichica, 7 6, 3 2, xii.19.48, H. B. Leech (CAS).
Puebla, 10 km. E., 5 ¢, 4 9, viii.24.62, JRZ (NMSU). Lago Totolzingo, Rt.
136, km. 200, 2 4, 7 2, viii.2.66, Flint & Ortiz (USNM). QUERETARO.
Queretaro, 1 ¢, 1 2, vi.20.57, D. R. Lauck (USNM); 16 6, 8 9, iti.27.63;
San Juan del Rio, 8 ¢, 7 2, ii.27.63, JRZ (NMSU). SAN LUIS POTOSI.

JAMES R. ZIMMERMAN 119

San Luis Potosi, 2m. S., 1 6, 1 2, xi.21.48, H. B. Leech (CAS). Santa Maria
del Rio, 5 6, 10 9, iii.26.63, JRZ (NMSU). SINALOA. Mazatlan, 7 m. S.,
1g, 1 9, xii.11.62, JIRZ (NMSU). VERACRUZ. Jalapa, 2 6, 2 2, Hoege
(ANSP); 4 6, 2 2, Hoege (AMNH). ZACATECAS. Fresnillo, 6600 feet,
16 m. NW., Rio Trujillo, 9 6, 14 2,, vi.29.54; 45 m. W., 7700 feet, 1 2,
vi.25.54, R. H. Brewer (CAS). Pinos, 10 m. S., 3 6,5 Q, viii.1.59; 5 m. S.,
4 8,1 9, viii.2.59, Ray Bandar (CAS). Sanchez Roman, 9 m. N., 5500 feet,
1 6,1 2, vii.17.54, R. H. Brewer (CAS).

Laccophilus mexicanus atristernalis Crotch, new status (Figs. 77, 80, 293)

Laccophilus atristernalis Crotch, 1873, p. 400; type not known; type locality
here restricted to Woodside, San Mateo County, California; Sharp, 1882a,
p. 292; Sharp, 1882b, p. 9; Horn, 1883, p. 277; Leng, 1920, p. 79; Zimmer-
mann, 1920, p. 22; Leech, 1948b, p. 401, Leech and Chandler, 1956, p. 314.

DIAGNOSIS. — This is the only irrorated Laccophilus in the northwestern
United States with black on the venter. Females may be brown, but males are
black. JL. biguttatus, a non-irrorated species with a black venter, does occur
rarely at higher elevations in the Sierras. All other sympatric Laccophilus are
brown or some lighter color beneath. A few females of L. m. decipiens and
atristernalis look so much alike that they are difficult to separate, but the former
is larger and has a more detailed elytral pattern.

VARIATION. — The most striking variation is the weakening of
the color on the venter of females from west of the Sierra Nevadas in
California and Oregon. While it is geographical, it is not racial.
Females of typical mexicanus from coastal southern California show
the same modification. Males of atristernalis and females from east
of the Sierras are as black beneath as any other races of mexicanus.
L. m. atristernalis may have a more suffused and reddish-brown ely-
tral pattern than mexicanus, also.

Males are larger than females (Table 5, Fig. 10) to about the
same degree as in mexicanus. The pronotal mean for males from
Churchill County, Nevada, is significantly different from the other
two samples, but the latter two do not differ from one another. More
samples are needed before assuming the difference is geographical
and not local ecological variation, however.

L. m. atristernalis and L. m. mexicanus are about the same size,
but the PW/EL ratio is different. The pronotum appears to be rela-
tively wider in atristernalis. If this proves to be the case, then it is
obvious that there is an important change in body proportions as well

MEM. AMER. ENT. SOC., 26

120 THE GENUS LaCCOPHILUS

as in aedeagus when atristernalis is compared to mexicanus. This
change serves as additional reason for maintaining them as separate
races.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— L. m. atristernalis ranges from southern Oregon and southwestern
Idaho to southern Nevada and California. Its eastern limit is central
Utah. It appears to be the most common Laccophilus in the Interior
Valley of California and around San Francisco Bay. It should inter-
grade with mexicanus in the southern part of California, Nevada, and
Utah. There is a single female from Victorville, San Bernadino
County, which has a dark brown venter and might be an intergrade
(Fig. 9). Anderson (1962) collected atristernalis in Utah below
5100 feet in small shallow bodies with little current and heavy sub-
mergent and emergent vegetation.

MATERIAL EXAMINED

UNITED STATES OF AMERICA.—CALIFORNIA. Alameda County.
Berkeley, 4, F. E. Winters (CNL); 4 ¢,5 9, F. E. Winters (CAS); 2 6, 2 2,
x.15.30 (BERK). Piedmont, 1 6, xii.4.11, F. W. Nunenmacher (FM). Butte
County. Oroville, 1 6, 1 2, v.29.21, R. C. Castleberry (AMNH). Colusa
County. Colusa, 1, v.31.35, E. C. Zimmerman (NMSU); 1 ¢, 1 92, vi.15.57,
T. R. Haig (DAV). Maxwell, 4, ix.5.18, A. Wetmore (USNM). Contra
Costa County. Mt. Diablo, 1 2, v.29.36, M. A. Embury (BERK). Fresno
County. Fresno, 2, iv.15.—, v.10.—, E. A. Schwarz (USNM); 1 2, vii.19.—
(AMNH). Selma, 3 6, vii.17.51, viii.12.51, viii.15.51, R. C. Bechtol & K. V.
Craig (DAV). Glenn County. Willows, 1 4, ix.8.49, M. Washbauer (BERK).
Inyo County. Deep Springs, 3 6, vii.17.53, W. D. McLellan (DAV); 1 2,
vi.19.54, P. Raven (CAS); 2 6, vii.15.53, J. R. Lattin (CAS). Keeler, 1,
vii.6.14 (USNM);1 2, 6 2, vii.6-14.00, Wickham (MCZ). Laws, 1 @, vii.26.38,
C. L. Hubbs (UMMZ). Olancha, 3400 feet, 6, G. R. Pilate (USNM). West-
gard Pass, 1 ?, vi.18.58, M. E. Irvin (DAV). Kern County. Bakersfield, 1 2,
v.5.31, A. T. McClay; 2 46, 3 2, vii.1.36, A. T. McClay (DAV); 1, iv.14.39
(BERK); 1 9, vii—.29, R. F. Smith (BERK). Elk Grove, 1 9, 1x.4.61,
J. K. Drew (CAS). Shafter, 20 m. W., light trap, 1 2, vi.18.56, 1 6, 4 9,
vii.20.56, T. R. Haig (DAV). Taft, 1, vii.5.37, H. M. Harris (ISU). Wasco,
3 2, vi.26.51, vi.27.51, vii.7.51, L. W. Isaak (DAV). County, 1 (USNM).
Kings County. Hanford, 7, R. S. Wagner (USNM). County, 13 ¢, 9 9,
vii—.33 (FM). Lassen County. Amadee, 1 6, 1 2, vii.21-28—, Wickham
(MCZ); 2, vii.21-28.— (USNM). Madera County. Cottonwood Ck., 3 ¢,
2 2, viii.26.50, P. S. Bartholomew (CAS). Northfork, 1, iii.19.20, H. Dietrich
(CNL); 1 6, vi.8.40, M. A. Cazier (AMNH). Marin County. County, 4,
x.18.19, H. Dietrich (CNL); 1 (USNM); ix.2.—, E. C. Van Dyke (AMNH).

JAMES R. ZIMMERMAN 121

Merced County. Athlone, 2 2, viii.l7— (AMNH). Los Banos, 2 6, 1 9,
iv.23.26, F. H. Wymore (DAV). Merced, 1 2, ix.—.42, W. F. Barr (FM).
San Joaquin R. at Hwy 152, 1 6, ii.2.64, H. B. Leech (CAS). Yosemite Lake,
1 6, x.8.31, R. Von (UMMZ). Modoc County. Cedarville, 5 m. E., 2 2,
vi.5.60, J. Schuh (SCH). Mono County. Benton, 1 ¢, vii.3.—, J. W. Tilden
(UMMZ). Mammoth, 1 6, 1 9, viii.28.46, G. P. MacKenzie (MCZ); 1 é,
1 Q, viii.28.46; 1 2, ix.20.45, G. P. MacKenzie (LACM). Nevada County.
Sagehen, 1 ¢, vii.21.51, R. C. Blaylock (DAV). Placer County. Auburn,
1 6, iv.11.51, A. T. McClay; Colfax, 4 6, v.11.21; Roseville, 1 6, v.13.46,
E. I. Schlinger (DAV). Sacramento County. Sacramento, 1 6, 3 2, v.20.44;
1 Q, vi.9.44;2 6,5 9, vii.2.44; 3 6, vii.5.44; 3 2, vii.9.44; 4 6, 1 2, vii.11.44;
2 2, vii.19.44; 2 6, vili.5.44; 1 2, vili.10.44; 3 2, vili.18.44; 1 36, ix.6.44,
A. T. McClay (DAV); 1, E. O. Essig (BERK). San Joaquin County. Banta,
4, xi.18.19, A. H. Hollonger (USNM). Stockton, 1, vii.8.08 (ARI). San
Mateo County. Woodside, 1 4, vi.23.52, P. S. Bartholomew (CAS). Shasta
County. Anderson, 1 2, vii.—.—, 1, viii.—.55, J. Willis (DAV). Hat Lake,
1 2, vi.2.47, E. E. Seibert; Lake Britton, 1 6, vi.29.47, C. H. Spitzer; Redding,
1 6,3 Q, vii.31.47, H. P. Chandler (CAS). Siskiyou County. Grenada, 1.3
m. E., 3 6,3 Q, viii.21.66, H. B. Leech (CAS). Solano County. Clarksburg,
2 2, vwii.10.31; 12 6, 17 9, vii.15.31; 7 2, vii.20.31; 1 2, viii.13.31, A. T.
McClay (DAV); 1, —.—.31 (BERK). Vacaville, 6 ¢, 6 2, vi.18.47; 1 6,
5 2, iv.15.47; 1 4, ix.10.47; 1, ix.21.47; 3 6, 1 9, ix.22.47;9 4, 6 Q, vii.13.48;
3 6, 7 Q, vil.14.48; 1 3, 72, vii.15.48; 16, 1 2, vii.26.48; 12 6, 12 9,
vii.27.48; 1 6, 2 9, vii.28.48; 1 3, 4 9, viii.6.48; 2 2, ix.6.48; 1 6, ix.10.48;
1 6, iv.8.49; 1 6, 2 9, vi.2.49; 1 9, viii.16.50, A. T. McClay (DAV). Stanis-
laus County. Del Puerto Canyon, 1 @, iii.22.46, W. F. Barr (FM). La
Grange, 2.8 m. E., Vizard Ck., 1 ¢, viii.19.62, H. B. Leech (CAS). Newman,
1 6, x.13.53, C. G. Moore (DAV). Sutter County. Robbins, 6, x.11.31,
H. P. Chandler (BERK). Yuba City, 1 ¢, v.10.42, H. P. Chandler (FM);
2, same data (BERK). Tehama County. Red Bluff, 1 ¢, vii.13.53, E. Yeo-
mann (DAY). Trinity County. Carrville, 2400-2500 feet, 1 2, v.25.34 (FM).
County, 1, v.23.34, T. H. G. Aitken (BERK). Tulare County. Hanford, 2 ?,
R. S. Wagner (CAS). Lindsay, 1 (CNL). Porterville, 1 ¢, viii.19.62, E. E.
Ball (SCH). Springville, 5 m. W., 3 ¢, 4 9, iv.15.54, R. K. Benjamin (USNM).
Three Rivers, 5 6, 4 2, Clberson (CNG). Visalia, 5 6,5 2, vii—.36, A. T.
McClay; 2 6, vili—36, A. T. McClay (DAV). Woodlake, 1 2, vi.24.36,
F. T. Scott (FM). County, 1 6, 1 2, vi.10.39, F. W. Nunenmacher (FM).
Yolo County. Davis, 4 4, ii.26.61, V. L. Vesterby; 1 ¢, iv.23.61, D. R. Miller
(DAV); 1 ¢, v.20.36, M. A. Embury; 2 2, v.—.36, J. J. Dubois (BERK); 2,
vii—.29 (BERK); 1 ¢, 1 2, v.22.25, F. H. Wymore; 1 2, x.—.46, A. L.
Booth; 1 6, 1 9, iv.19.47, B. Stevens; 1 6, v.15.47, E. I. Schlinger; 5 $, 4 9,
Vii.27.48; 1 6, 1 2, vii.30.48; 2 2, vili.1.48; 1 ¢, 2 2, ix.9.48; 1 3, iii.31.50,
A. T. McClay; 6 ¢, 3 2, viii.19.50; 3 6, 12 9, viii.13.51, R. C. Bechtol; 1 6,
12> 1v-25.58,, Po Raige; 1 9,1x-29.59, M. B: Irwin; 1 6; 3 2, x-15.60) D: OQ:
Cavagnaro; 1 6, 1 2, ix.24.60, D. Q. Cavagnaro (DAV). Dixon, 1 2, ix.14.47,

MEM. AMER. ENT. SOC., 26

122 THE GENUS LACCOPHILUS

A. T. McClay (DAV). Elkhorn Ferry, 1 ¢, vii.9.51, E. I. Schlinger (DAV).
Monticello, Putah Ck., 9, xiii—.17, J. C. Bradley (CNL). Rumsey, 1 ¢,
vii.23.55, E. A. Kurtz (DAV). Woodland, 1 6, 1 92, vii.4.46 (BERK); 1 9,
vii.4.47, E. G. Meyers (BERK). IDAHO. Owyhoe County. Hot Springs,
1 4, 2 9, vii.17.49; 1 2, vi.28.53, W. F. Barr (FM). NEVADA. Churchill
County. Fallon, 14 6, 19 9, ix.2.59, F. D. Parker (DAV). Elko County.
Ruby Valley, hot springs (65 m. S. Wells), 2 6, 2 2 (MCZ). Lander County.
Battle Mtn., 37 m. S., hot springs, 1 2 (MCZ). Lincoln County. Maynard
Lake, 3 6, 4 9, vii.12.38, Calhoun (CAS). Mineral County. Hawthorne,
4567 feet, 1, vii.27.— (USNM). Walker Lake, 1 ¢, xii.29.38, F. D. Parker
(DAV). Nye County. Amargosa Hot Springs, 1 ¢, 3 9, iii.31.53, H. B.
Leech (CAS). Ash Meadow Lodge (near Rhyolite), 10 6, 3 2, vii.10.54,
N. L. Rump (AMNH); 3 2, iii.30.66, J. Schuh (SCH). Beatty, 3 m. N.
Amargosa R., 1 6, 1 9, iii.29.66; 2 m. N., 2 6, 1 9, i1i.29.66, J. Schuh (SCH).
Springdale, 2, vi.11.36, J. G. Needham (CNL). Washoe County. Reno, 3,
iii.6.40, J. G. Needham (CNL); 2 6, 2 2? (AMNH); 1 4, Wickham (ANSP);
1 6,1 9, vi.3.40, La Rivers (MCZ); 1 6, v.23.41, I. La Rivers (CAS); 1 ¢,
vi.3.40 (CAS). OREGON. Harney County. Crane Hot Springs, 25 m. SE.,
Burns, 4 6, 3 9, ii.25.65, K. Goedon (ORES). Fields, 5 m. NE., 1 2, vi.4.63,
K. Goedon (ORES). Harney Lake, 0.5 m. S., 3 ¢, 3 2, vi.5.63, K. Goedon
(ORES). Jackson County. Medford, 1 ¢, viii.29.44, A. T. McClay (DAV);
6m. N., 1 4, 1 2, v.22.60, J. Schuh (SCH). Lake County. Albert Lake,
1 2, vi.6.58; 6 6, 3 9, vi.7.58, Vertrees & Schuh (SCH). UTAH. Salt Lake
County. Salt Lake, 1 9, ix.23.57, R. D. Anderson (RDA). Tooele County.
Timpie, 1 m. S., 1 2, v.1.57, R. D. Anderson (RDA). Utah County. Salem
Pond, 4000 feet, 1 6, vii.17.41, H. P. Chandler (FM). Utah Lake, 1 4,
vi.19.— (ANSP).

Laccophilus mexicanus oaxacensis, new subspecies
(Figs. 74, 78, 79, 295)

DIAGNOSIS AND DESCRIPTION. — This race differs from L. m. mexi-
canus and L. m. atristernalis in its larger size (Table 5, fig. 10), more detailed
elytral pattern, more rounded female last ventral segment, more elongated aede-
agus, and in the comparatively longer male fifth pro- and mesotarsal segments.
The elytral pattern has the anterior fingerlike projections that are present in so
many Laccophilus, but in neither of the other two races of mexicanus. Lateral
clear areas are also more distinct, and there are frequently hollowed elongate
marks on either side of the midline at about the level of the midlateral maculae.
The epipleural flange, when it is present, begins just behind the midlateral macula.
The male coxal file has 25 to 30 easily counted lines and about 15 much finer
and less distinct ones. There is a weak file in females. The male last ventral
segment has fewer rugae than mexicanus, and the female segment has depres-
sions on either side of the median crest with protuberances anteriolateral to the
depressions. The fifth protarsal segment is twice as long as the fourth; the

JAMES R. ZIMMERMAN 123

fifth mesotarsal one is one and seven-eighths as long as the fourth.

L. m. oaxacensis can be separated from the sympatric L. mistecus mistecus
by its smaller size, different aedeagus, and the rounded (rather than slightly
truncated) last ventral segment.

30 4
oaxacensis

o.

Z

"A
2,

7)

mexicanus

KN) WW IM) th fe ig 8

4.0 4.2 4.4 4.6 48 5.0 5.2 5.4

THOME AEE EONiGaiy He (mim)
Figure. 10. Histograms of length of Laccophilus mexicanus. Males are
shown crosshatched; females, stippled.

VARIATION. — The range in length varies from about 4.32
mm in the smallest female to about 5.40 mm in the largest male.
Average lengths vary from 4.79 mm in the smallest female sample to

MEM. AMER. ENT. SOC., 26

124 THE GENUS LACCOPHILUS

5.14 mm in the largest male sample (Table 5). Males are usually
larger than females with a peak frequency at about 5.10 mm.

The most common variation in pattern is the degree to which the
elytral pattern is margined or the occurrence and intensity of the
mark that occurs on the disc near the midline. The amount of dark-
ening on the abdominal sclerites is also variable.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This race has been collected only within fifty miles of the valley
of Oaxaca at elevations from about 5000 to 7000 feet. It is one of
the most common aquatic beetles in that area, however. It was usual-
ly taken in roadside ditches and shallow quarries, but occurs more
infrequently in pools of grassy margined streams. It appears to pre-
fer habitats with mud bottoms.

MATERIAL EXAMINED

Holotype male, allotype female; and five male and five female paratypes
from three miles north of Oaxaca, Oaxaca, vili.25.63, J. R. Zimmerman, are
deposited in the United States National Museum. Other paratypes deposited
as follows: seven males and 19 females from eight miles north of Oaxaca,
Oaxaca, xii.12.48, H. B. Leech, are in the California Academy of Science, San
Francisco; a male and female also from the preceding locality are in the Uni-
versity of Michigan Museum of Zoology, Ann Arbor; two males and two fe-
males with the same locality data as the holotype are also in the University of
Michigan Museum of Zoology and in the Departmento de Entomologia, Labora-
torio de Sanidad Vegetal, Coyoacan, D. F., Mexico.

Laccophilus pseudomexicanus, new species (Figs. 82-89, 296)

DIAGNOSIS. — This species belongs to the group with an irrorated pattern,
metacoxal file, sawlike ovipositor, and has black on the venter. It is larger
than mexicanus and salvini, the same size as spergatus, and is smaller than
mistecus. It overlaps all of those, but salvini, however. Reliable separations
require comparison of the aedeagi, but the pronotum is relatively longer in
pseudomexicanus (LP/PW, 0.43) than in any of the others (0.40-0.41); and
the males have the pro- and mesotarsi enlarged in the lateral as well as dorso-
ventral plane.

DESCRIPTION. — Large (length, 4.7 to 5.7 mm; width, 2.5 to 3.3 mm),
brown, irrorated species; black on venter; metacoxal file present; prosternal
process short; sawlike ovipositor. COLOR. Head: light yellowish-brown dark-
ening to brown or reddish-brown on occiput near the pronotum; appendages
yellow except mandibles which darken toward tip. Pronotum: light yellowish-
brown; disc frequently with obscure darker paired marks; posterior margin and
apex may be darker due to translucence rather than additional pigment. Elytra:

JAMES R. ZIMMERMAN 125

strongly irrorate with brown over almost complete surface, but midlateral spot
present. Tergite VIII: dark brown to black. Venter: prosternum, its process,
fore and middle legs light brownish-yellow with reddish tinge; hind legs of
about the same color, but with much stronger reddish highlights; mesosternum,
metasternum, metacoxal plates, except postcoxal processes, very dark brown or
black; abdominal segments frequently pale except for the black hind margin of
first visible segment; other segments often suffused with dark brown or black.
Genitalia: oval plate, aedeagus, and parameres with varying mounts of dark
reddish-brown or yellowish-brown; ovipositor reddish-brown. ANATOMY.
Microreticulation: weakly double on head, pronotum, and elytra; cellules small
and uniform. Head: supraclypeal seam closely parallel to margin. Pronotum:
WH/PW, 0.68; LP/PW, 0.43. Elytra: large epipleural flange in some females;
apical truncation marked. Venter: coxal file with about 25 to 30 fine lines which
grade into the microsculpture of the metacoxae; prosternal process with well-
defined crest; lobes of postcoxal laterally projecting well beyond midline; last
visible ventral abdominal segment of male somewhat truncate, produced and
with an asymmetrical crest nearly attaining the apex; a few small rugae along
the posterior margin; female last ventral segment not truncated; median crest
poorly defined, but several rugose-like lines converging toward midline near the
apex; oblique striations present and strongly impressed; posterior margin re-
flexed; an oblique low ridge which does not attain the apex present on either
side of median crest; setigerous punctures spread over surface, but thickest at
middle and near the posterior margin. Legs: male protarsi expanded laterally
as much as in the dorsoventral plane; mesotarsi expanded more dorsoventrally
than laterally; palettes easily seen at 20 power magnification; male fifth tarsal
segment about one and a half times as long as fourth in both pair of front legs;
profemoral setae (5 to 9) margin finer and shorter than mesofemoral ones (6).
Genitalia: oval plate large with sharp, slightly produced tip; median crest strong
apically, but weakened to raised line anteriorly; raised lines on right side of
crest diverge to right anteriorly; aedeagus bent only slightly, but with distinctive
flattened knob at apex; ovipositor with 15 to 16 sawlike teeth.

VARIATION. — Males are about 0.2 mm larger than females
(Table 6, Fig. 11). The sample from San Luis Potosi suggested
that there is a racial difference between it and the population in
Durango and Jalisco. The populations from the latter two areas, on
the other hand, appear to be very similar. Elytral pattern differences
support the same conclusion. Specimens from San Luis Potosi have
local concentration of pigment on the elytral disc, while the others
have a more uniform irroration.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This species is known only from the Mexican highlands in Durango,
San Luis Potosi, Michoacan, and Mexico. It is found in both the

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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JAMES R. ZIMMERMAN 127

Sierra Oriental and Sierra Occidental and in the high valleys of the
state of Mexico. I have collected it mainly in mountain stream pools
with bottom of mud and volcanic gravel.

MATERIAL EXAMINED

Holotype male, allotype, and two paratypes of each sex with the following
data are in the United States National Museum; 5 km. E. of Mazamitla, Jalisco,
i1i.27.64, J. R. Zimmerman. Other paratypes deposited as follows: seven males
and three females from 15 miles east of Cuidad del Maiz, San Luis Potosi,
xi.19.48, H. B. Leech, and one male from 15 to 20 miles west of Jiquilpan,
Michoacan, xi.30.48, E. S. Ross, are in the California Academy of Sciences,
San Francisco; one male from 8.7 miles NW. of Acambay, Mexico, 8300 feet,
ii.3.53, I. J. Cantrell, and one female with the same data as the holotype is in
the University of Michigan Museum of Zoology, Ann Arbor; and one male and
one female with the same data as the holotype are also in the Departmento de
Entomologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F., Mexico.

MEXICO. — DURANGO. Llano Grande, 10 m. E. (Navios), 3 4, 6 2,
xii.10.62; 3 m. E., 1 6, xii.10.62, JRZ (NMSU). El Salto, 16 m. E., 16 4,
21 2, vi.29.64, P. J. Spangler (USNM). JALISCO. Ameca, 25 km. E.,
1 2, iii.25.64; Mazamitla, 5 km. E., 22 ¢, 27 9, iii.27.64. Near Tala, 1 é,
2 2, ili.25.64; near Tamazula, Rd. 110, 1 2, ili.27.64; Zapotlenejo, 7 m. E.,
6 6,62, JRZ (NMSU). MEXICO. Acambay, 1 ¢, 5 2, iii.27.63; Atloco-
mulco, 1 6, 1 Q, iii.27.63, JRZ (NMSU). Ixtlahuaca, 6 m. S., 2 2, vii.12.52
(53?), R. & J. Selander (USNM). MICHOACAN. Jiquilpan, 8 km. E., 2 ¢,
ii.27.64, JRZ (NMSU). SAN LUIS POTOSI. Agua Zarca (near Platinito),
5 6,2 Q, iii.25.63, IRZ (NMSU).

LACCOPHILUS MISTECUS
(Figs. 90-97, 298)

This species is composed of two races whose ranges are too in-
completely known to be able to give more than a rough approxima-
tion of their distribution. They are both confined to Mexico and at
the moment seem to be restricted to north of the Isthmus of Tehuan-
tepec. The races approach one another in adjacent areas of Jalisco
and Michoacan. There is a large collecting gap from Oaxaca through
Guerrero and Michoacan. The principal difference between the two
races is that the more northern one is light beneath and southern one
is black or dark brown beneath. The species description is given
under L. m. aztecus, new subspecies.

NOMENCLATURAL NOTES. — This species has several dis-
tinctive anatomical features and a limited distribution; consequently,

MEM. AMER. ENT. SOC., 26

128 THE GENUS LACCOPHILUS

no nomenclatural confusion has developed. Sharp gives Parada as
the type locality for this species. That locality is La Parada, Oaxaca,
Mexico, which Selander and Vaurie (1962) locate as a “Hacienda
and important collecting site on the north slope of the mountains west
of Cerro San Felipe, which is just north of the city of Oaxaca; 7900
feet, about 17° 10’, 96° 40’.”. This means that the dark ventered
race is the nominate one and that the light colored one should be
described as new.

Laccophilus mistecus mistecus Sharp, new status

Laccophilus mistecus Sharp, 1882b, p. 9. Holotype: male, Parada, Mexico,
British Museum (Natural History) (Salle Coll.); Zimmermann, 1920, p. 23;
Blackwelder, 1944, p. 74.

DIAGNOSIS. — This is the largest of the species which have a metacoxal
file, are irrorated brown above, and have black beneath. The truncate last
ventral segment with its prominent lobe in the male is also a distinctive feature.
L. mistecus resembles spergatus in this character; but there is some local ac-
cumulation of pigment on the elytra, in contrast to the evenly irrorated pattern
of spergatus. The ventral crest on the oval plate of m. mistecus is much less
prominent than in spergatus, also. L. pseudomexicanus has the last ventral seg-
ment untruncated in the male. Male aedeagi will separate all three of these
species. L. mexicanus and L. salvini are much smaller with few individuals
exceeding 5.4 mm. The description given for L. mistecus aztecus can be used
for L. m. mistecus except, in the latter, the venter has black on it instead of
paler shades of brown.

VARIATION. — Males are 0.25 mm to 0.30 mm larger than
females (Table 8, Fig. 11). Males average just over 6.0 mm long
and females just under 6.0 mm. The quantitative data conflicts with
the separation into two races on the basis of the venter color. The
sample from Michoacan more closely resembles the Acambay, Mex-
ico, sample than the one from Oaxaca.

MATERIAL EXAMINED

MEXICO. — JALISCO. Mazamitla, 5 km. E., 2 6, iti.27.64; Tamazula (ject.
Rd. 110), 2 9, iii.27.64, JRZ (NMSU). MICHOACAN. Jiquilpan, 11 m. W.,
9 3, 6 @, vii.28.62, IRZ (NMSU). OAXACA. Nochixtlan, 2 km. S., 2 2,
ix.4.64; Ixtlan de Juarez, 32 6, 50 9, viii.25.63, JRZ (NMSU). Tutla, 3 3,
1 9, xii.13.48, H. B. Leech (CAS).

JAMES R. ZIMMERMAN 129

pseudomexicanus

spergatus

NUMBERS

mistecus mistecus

4.8 5.0 5.2 5.4 5.6 5.8 : 6.2

10
mistecus aztecus
O aes

4.8 5.0 5.2 5.4 5.6 5.8 6.0 6.2

YO wat LIEN © wl (non)

Figure 11. Histograms of length in Laccophilus pseudomexicanus, L. sper-
gatus, L. mistecus. Males are shown crosshatched; females, stippled.

Laccophilus mistecus aztecus, new subspecies

DIAGNOSIS. — This differs from L. m. mistecus only in the presence of
a pale venter. It is most likely confused with L. maculosus shermani which
also occurs in central Mexico. They can be separated by elytral pattern details,
rounded male segment in shermani, and the presence of an epipleural flange in
females of the latter.

MEM. AMER. ENT. SOC., 26

130 THE GENUS LACCOPHILUS

DESCRIPTION. — Large (length, 5.4 to 6.3 mm; width, 3.2 to 3.7 mm),
brown, irrorated subspecies; metacoxal file present; prosternal process short;
ovipositor sawlike. COLOR. Head: light yellowish-brown above and beneath;
darker at pronotum between the eyes; appendages yellow except mandibles
which are reddish-brown; gular sutures dark brown. Pronotum: light yellowish-
brown with slight darkening in the posterior half of the disc. Elytra: fairly
evenly irrorated with brown on a light yellowish-brown background, but with
some suffusion and coalescence of dots; epipleura pale anteriorly and dark
posteriorly. Tergite VIII: dark brown to black. Venter: yellowish-brown.
Genitalia: variably yellowish-brown to reddish-brown to black. ANATOMY.
Microreticulation: weakly double on head, pronotum, and elytra. Head: supra-
clypeal seam closely parallel to margin. Pronotum: WH/PW, 0.67; LP/PW,
0.40. Elytra: apical truncation slight; epipleura flange rarely, if ever, present.
Venter: coxal file prominent in males, composed of 35 to 40 lines; prosternal
process with well-defined crest; lobes of postcoxal processes rounded and later-
ally projecting posterior to midline; last male ventral abdominal segment trun-
cated and produced, but less so than in spergatus; asymmetrical median crest
and anteriolateral crests present, forming concavities on either side, the left
larger than the right; corresponding female segment not truncated, but pyrimidal
with a groove on either side forming an emarginate edge; setigerous punctures
scattered over entire surface, but thickest near apex; crest broad and triangular,
not ridgelike. Legs: male pro- and mesotarsi slightly enlarged in a dorsoventral
plane; palettes easily observable at 20 power magnification; fifth protarsal seg-
ment not quite twice as long as fourth; fifth mesotarsal segment one and two-
thirds as long as fourth; profemoral setae (5 to 6) shorter and finer than
mesofemoral ones (5 to 7, usually 7). Genitalia: oval plate large with acumi-
nate tip; its median crest curving slightly to the left and extending nearly to
anterior margin; about twice as many lines on the right side of the crest as on
the left; right paramere with blunt tip; left with smoothly rounded apex; aede-
agus with cuneate projection about half the distance to the apex and one at the
apex; Ovipositor with about 15 sawlike teeth.

MATERIAL EXAMINED

Holotype male, allotype, and a male and female with the following data
are in the United States National Museum; Acambay, Mexico, iii.27.63, J. R.
Zimmerman. Other paratypes distributed as follows: one male, two females,
12 m. S. of Ojuelos de Jalisco, Jalisco, xi.21.48, H. B. Leech; one male, 10 m.
S. Pinos, Zacatecas, vili.1.59, Ray Bandar, California Academy of Sciences, San
Francisco, California; one male, 8.7 m. NNW. of Acambay, Mexico, 8300 feet,
i1.3.53, I. J. Cantrell, and a female with the same data as the holotype, Uni-
versity of Michigan Museum of Zoology, Ann Arbor, Michigan; one male and
one female with the same data as holotype, Departmento de Entomologia,
Laboratorio de Vegetal Sanidad, Coyoacan, D. F. Mexico.

MEXICO. — JALISCO. Ojuelos de Jalisco, 12 m. S., 1 ¢, 2 9, xi.21.48, H. B.
Leech (CAS). MEXICO. Acambay, 3 6,5 2, iti.27.63, JRZ (NMSU); 8.7

131

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MEM. AMER. ENT. SOC., 26

132 THE GENUS LACCOPHILUS

m. NNW., 8300 feet, 1 ¢, 11.3.53, I. J. Cantrell (UMMZ). Atlocomulco, 2 6,
5 9, iii.27.63, JIRZ (NMSU). MICHOACAN. Quiroga, 1 6, 2 9, vili.10.57,
D. Lauck (USNM). SAN LUIS POTOSI. S. Maria del Rio, 1 9, iii.26.63,
JRZ (NMSU). DURANGO. Llano Grande, 10 m. E. (Navios), 3 6, 6 2,
xii.10.62; 3 m. E., 1 6, xii.10.62, JRZ (NMSU). El Salto, 16 m. E., 16
21 2, vi.29.64, P. J. Spangler (USNM). JALISCO. Ameca, 25 km. E., 1
11.25.64; Mazamitla, 5 km. E., 22 6, 23 9, iii.27.64. Near Tala, 1 6, 2
11.25.64; near Tamazula, Rd. 110, 1 9, 11i1.27.64; Zapotlenejo, 7 m. E., 6 ¢, 6

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1 @, iii.27.63, JRZ (NMSU). Ixtlahuaca, 6 m. S., 2 9, vii.12.52 (537), R. &
J. Selander (USNM). MICHOACAN. Jiquilpan, 8 km. E., 2 ¢, iii.27.64,
JRZ (NMSU). SAN LUIS POTOSI. Agua Zarca (near Platinito) 5 ¢, 2 9,
11.25.63, JRZ (NMSU).

LACCOPHILUS SPERGATUS e

:

LACCOPHILUS M. MISTECUS ®&
LACCOPHILUS M AZTECUS a

ACCOPHILUS FUSCIPENNIS o

Figure 12. Distribution of Laccophilus mistecus, L. spergatus, and L. fusci-
pennis.

JAMES R. ZIMMERMAN 133

Laccophilus spergatus Sharp (Figs. 98-105, 297)

Laccophilus spergatus Sharp, 1882b, p. 10. Holotype: female, Toluca, Mexico,
British Museum (Natural History); Zimmermann, 1920, p. 26; Blackwelder,
1944, p. 74.

DIAGNOSIS. — Size, the remarkably evenly irrorate elytra, black venter,
and strongly truncate last ventral segment in males and females separate this
species from most other North American Laccophilus. Examination of the
distinctive male genitalia may be necessary to separate it from some specimens
of mistecus, pseudomexicanus, or mexicanus, however. The truncated female
ventral segment of spergatus separates them from females of the last two spe-
cies; but the male ventral segment of mistecus is truncated, and an examination
of the aedeagus probably is necessary.

DESCRIPTION. — Large (length, 5.08 to 6.16 mm; width, 2.92 to 3.46
mm), brown evenly irrorated species; black beneath; metacoxal file present;
prosternal process short; ovipositor rakelike. COLOR. Head: brownish-yellow
above and beneath darkening to light brown at base of pronotum between the
eyes; appendages yellow, except mandibles which are reddish-brown at tips;
gular margin nearly black. Pronotum: pale brownish-yellow with slightly
darker tinge on the disc. Elytra: irroration uniform over almost entire elytra
and, while some of the dots occur in strings, there is little or no coalescence of
dots nor suffusion of color between dots; in males some weakening of the irrora-
tion at humeral angle and along the sutural margin in the posterior half; epi-
pleura pale anteriorly and dark reddish-brown posteriorly. Tergite VIII: dark
brown to black. Venter: prothorax, its process, pro- fore and mesolegs all pale
brownish-yellow; mesothorax, metathorax, and metacoxal plates dark brown to
black except for lighter postcoxal processes; abdominal sternites generally pale
yellow-brown to light reddish-brown, seldom black, except for the suture be-
tween the first and second segment; oblique striations usually darkened and
prominent. Genitalia: variably pale yellow-brown to dark reddish-brown; usu-
ally evenly pigmented. ANATOMY. Méicroreticulation: faintly double, cellules
more deeply impressed in females than males; hence the latter more shining.
Head: supraclypeal seam closely parallel to margin. Pronotum: WH/PW,
0.68; LP/PW, 0.40. Elytra: epipleural flange absent; apical truncation slight,
but more apparent in females than males. Venter: coxal file prominent in
males (33 to 36 lines) and faintly suggested in females; prosternal process with
crest well-defined in apical half; postcoxal processes rounded and laterally pro-
jecting beyond midline; last visible ventral abdominal segment of male sharply
truncated with produced apex; median crest bent sharply to the left; strong
ridges extending to the anterior segment are near the lateral margins; the larger
left one with a marked depression situated posteriorly and medially; female seg-
ment less truncated than male and without definite median crest; apex flattened.
Legs: male pro- and mesotarsi enlarged dorsoventrally and produced on either
side; palettes easily visible at 20 power magnification; fifth tarsal segment of
pro- and mesotarsi slightly less than twice as long as the corresponding fourth

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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segment; profemoral setae (5 to 7) shorter and finer than mesofemoral ones
(5S to 7). Genitalia: oval plate and aedeagus are the most distinctive of the
genus in North America; oval plate with rounded apex and small tip; large
ventral keel bending to right anteriorly; several other large ridges to the right of
the keel; aedeagus with two large lateral projections, one placed at about half
the length and the other at the apex; right paramere with a produced slightly
curved tip; ovipositor with about 16 sawlike teeth.

NOMENCLATURAL NOTES. — Sharp described this species
from two female specimens; but fortunately, the female is sufficiently
distinct from related species that identification and separation have
good reliability. A precise locality was also given; and I have col-
lected and examined numerous specimens from near Toluca, the type
locality. I have examined the cotype which is from that same local-
ity. F.N. Young has also sent me a drawing and notes on the type.

VARIATION. — Males are about 0.2 mm larger than females
(Table 2, Fig. 11). The largest male and female measures 6.15 mm
and 5.79 mm respectively. The means for lengths were about 5.78
mm and 5.57 mm. L. spergatus is relatively narrower than most ir-
rorated species, and this is reflected in the WP/EL ratio (0.543 to
0.549). The difference in means in pronotal width between males
from Mexico and Durango is significant by “t” test at the ninety per-
cent level. Specimens from all localities resemble one another closely,
however; and there do not appear to be any racial differences.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This species has a distribution limited to above 5000 feet from
Durango to the state of Mexico. Collections were made from between
6000 to 9000 feet in small pools and sluggish streams or in mountain
meadow seepage areas with fine, gravelly soils and grassy margins.

MATERIAL EXAMINED

MEXICO. — DURANGO. Durango, 20 m. W., 7500 feet, 7 4, 12 2, xii.10.62,
JRZ (NMSU); 25 m. W., 20+, vi.29.64, P. J. Spangler (USNM). El Pino,
5 m.N., 1 9, vii.7.52, J. D. Lattin (BERK). El Salto, 16 m. E., 14 6, 16 9,
vi.29.64, P. J. Spangler (USNM). Las Adjuntas, 2 m E., 2 é, 2 9, vi.30.52,
J. D. Lattin (BERK). Llano Grande, 10 m. E. (Navios), 3 6, 23 2, xii.10.62;
3m. E., 7 6, 25 2, xii.10.62, JRZ (NMSU). MEXICO. Acambay, 8.7 m.
NNW., 8300 feet, 1 6, 2 9, ii.3.53, I. J. Cantrall (UMMZ); 23 6, 23 9,
11.27.63; near Atlahuaca, 10 9, 13 9, viii.29.62; Atlocomulco, 7 é, 8 Q,
11.27.63, JRZ (NMSU). La Marquesa, 1 m. W., 4 6, x11.8.48, H. B. Leech
(CAS). QUERETARO. San Juan del Rio, 1 6, 1 2, iii.27.63, JRZ (NMSU).

MEM. AMER. ENT. SOC., 26

136 THE GENUS LACCOPHILUS

SAN LUIS POTOSI. San Luis Potosi, 2 m. S., 2 6,1 2, xi.21.48, H. B. Leech
(CAS). Santa Maria del Rio, 1 ¢, iii.26.63, JIRZ (NMSU). ZACATECAS.
Fresnillo, 61 m. W., 8100 feet, 1 6, 4 9, vi.25.54, R. H. Brewer (CAS).

Laccophilus fuscipennis Sharp (Figs. 106-113, 290)

Laccophilus fuscipennis Sharp, 1882b, p. 10. Holotype: female, Oaxaca, Mex-
ico, Hoege, British Museum (Natural History); Zimmermann, 1920, p. 19;
Blackwelder, 1944, p. 74.

DIAGNOSIS. — This is a medium-sized species which belongs to the group
with brown irrorated elytra, pale venter, metacoxal file, short prosternal proc-
ess, and sawlike ovipositor. It is sympatric with similar species that share some
or all of those characters. It has several distinctive secondary sexual charac-
ters, however. It is dorsally similar to mexicanus, salvini, proximus, spangleri,
and vacaensis. It differs from the first two in that they are dark beneath.
L. proximus, L. spangleri, and L. vacaensis have considerably more pattern on
the elytra than fuscipennis (figs. 289, 312-315). L. fuscipennis has a meta-
coxal file in the male and a sawlike ovipositor in the female, while spangleri
and vacaensis lack files and have forklike ovipositors. It may be necessary, if
pattern details are obscured, to compare males of proximus (confusus Sharp)
and fuscipennis, which have quite different aedeagi.

DESCRIPTION. — Small or medium size (length, 3.9 to 4.7 mm; width,
2.3 to 2.7 mm), light brown, irrorated species, pale beneath; metacoxal file pres-
ent; prosternal process short; ovipositor sawlike. COLOR. Head: yellowish-
brown, with some darkening between the eyes at the base of the pronotum.
Pronotum: yellowish-brown. Elytra: pale brownish-yellow background, irrora-
tion relatively uniform over the anterior half of the elytra, but considerable
suffusion and darkening of color between irrorations in the posterior half; pat-
tern margins poorly defined; apex usually without darker pigment; anterior half
of epipleura pale; posterior half may be darkened or pale. Tergite VIII: dark
brown. Venter: pale yellowish-brown with some reddish-brown highlights
around the mesocoxae, the postcoxal processes, and hind legs. Genitalia: oval
plate, bases of parameres, base and tip of aedeagus reddish-brown; parts of
parameres and shaft of aedeagus reddish-yellow-brown. ANATOMY. Micro-
reticulation: weakly double on head, pronotum and elytra. Head: supra-
clypeal seam arching upward at midline. Pronotum: WH/PW, 0.70; LP/PW,
0.40. Elytra: epipleural flange in about 50 per cent of all females; expansion
starting at about midbody level, i.e., at the front of the position of the mid-
lateral spot; apices slightly truncated. Venter: coxal file only weakly present
in males; ridges counted with difficulty (about 25 to 30); prosternal process
with well-defined crest; lobes of postcoxal processes rounded and laterally pro-
jecting well beyond the midline; male right posterior margin of fifth abdominal
segment with slight protuberance; last ventral abdominal segment of male slightly
truncated, produced with rugae along the posterior margin; margin of female
last ventral segment weakly sinuate on either side and with a distinct groove
forming a marginate edge; midline crest well-developed with a nearly acute

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AMER. ENT. SOC.,

MEM.

138 THE GENUS LACCOPHILUS

peak at apex; a few rugae near apex and at middle of each side. Legs: male
pro- and mesotarsi noticeably enlarged in dorsoventral plane; palettes large
enough to be easily distinguished at 20 power magnification, fifth tarsal segment
on both pair of legs about one and four-fifths as long as corresponding fourth;
5 to 6 setae on both pro- and mesofemoral margins. Genitalia: oval plate
large, with sharp, but not acuminate tip and numerous (20 to 25) raised lines,
but with weak crest apparent only near apex; crest and lines all strongly curved
to the left; aedeagus with distinctive apical cuneiform projection, bent with
groove on one side; right paramere elongated with produced apex; left para-
mere with rather broad rounded base; ovipositor with about 12 sawlike teeth.

NOMENCLATURAL NOTES. — There appears to be no syn-
onymy in this species, even though Sharp described it from just two
females and it has a fairly wide distribution over Mexico. The type
locality is Oaxaca, Mexico. I have not been able to collect this
species from near Oaxaca City, but I have taken it at a lower eleva-
tion on the Isthmus of Tehuantepec. Apparently, some males from
other species were placed with the two females in the collection of the
British Museum (Natural History); but Sharp clearly marked a type.
I have seen the cotype, which appears to be practically identical with the
type, and have compared the cotype with numerous other specimens.

VARIATION. — Some individuals have less uniform irroration
than others, with darker pigment accumulating in the posterior half
of the elytra in a manner suggestive of salvini or mexicanus; but the
variation does not seem to have a geographical basis.

Females from the west coast generally have smaller epipleural
flanges than do those from Veracruz. The incidence of females with
flanges is just as high, however.

The sexes are about the same size. The peaks in length frequency
distributions are at the same measurement — about 4.75 mm. The
overlap is nearly complete, but the largest individuals were males and
the smallest ones were females. Populations from both coasts agree
closely in quantitative characters and give no indication of any racial
difference (Fig. 16).

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This is a Mexican species that occurs commonly on the coastal
lowlands and rarely at higher elevations. It has been taken from all
across the Isthmus and northwestern Chiapas to San Luis Potosi and
on the west coast from the southern coast of Jalisco to Mazatlan.
There is a large gap between east and west coast populations, un-
doubtedly due to a lack of collections. It occurs mainly in temporary

JAMES R. ZIMMERMAN 139

situations in clay soils. I have collected it most frequently in road-
side ditches and cattle ponds that have little vegetation and few rocks
or gravel. It is clearly a tropical species that has its northern limits
restricted by freezing temperature (Fig. 12).

MATERIAL EXAMINED

MEXICO. — CHIAPAS. Near Pichucalco, 1 ¢, 1 2, xii.26.63, JRZ (NMSU).
CHIHUAHUA. Hidalgo del Parral, 1 2, xii.9.62, JRZ (NMSU). JALISCO.
Barra de Navidad, 1 ¢, x.25.66; La Huerta, 5m. N., 1 ¢, x.25.66, A. H. Smith
& JRZ; Mezquitan, 3 6, 1 Q, iii.28.64, JRZ (NMSU). MORELOS. Cuerna-
vaca, 1 2, vili—.55, L. H. Krauss (CAS). NAYARIT. Acaponeta, 8 m.
NW., Rio de las Canyas, 6 ¢6, 2 92, xi.25.48, H. B. Leech (CAS); 12 m. S.,
1 Q, vii.31.62, JRZ (NMSU). Pena, 12 m. N., 1 4, 3 9, xi.26.48, H. B.
Leech (CAS). San Blas, 5 m. E., 3 ¢, 1 2, vii.31.62, JRZ (NMSU). Tepic,
18 m. NW., 1 6, xi.27.48, H. B. Leech (CAS). OAXACA. Matias Romero,
2 2, ix.9.64; Juchitan, 20 km. N., 1 6, 1 2, ix.7.64, JRZ (NMSU). PUEBLA.
Near Maria Andrea, 12 4, 19 92, ix.10.64, JRZ (NMSU). SAN LUIS POTOSI.
Comoca, Rio Axtla, Rd. 85, 7 6, 4 9, ili.23.63, JRZ (NMSU). Cuidad del
Maiz, 1 2, xi.19.48, H. B. Leech (CAS). SINALOA. Concordia, 8 m. E.,
2B, OS sei lAOwre IMevanienn, WG 8, WS Vs oatoiilwe 7/ ie Sey Wh Sy Ite
xii.11.62, JRZ (NMSU). VERACRUZ. Acayucan, 20 m. S., 10 ¢, 11 2,
ix.7.64; Catemaco, 1 ¢, 1 9, viii.26.62, JRZ (NMSU). Cuitlahuac, 20 2,
20 @, viii.10-12.64, P. J. Spangler (USNM). WHuatusco, 25 km. E., 1 6, 3 2,
ix.9.64, JRZ (NMSU). Near Garro, 13 ¢, 10 2, ix.8.64: J. D. Covarrubia,
1 6, 4 &, vili.26.62; Martinez de la Torre, 5 km. E., 5 6, 9 2, ix.9.64; near
La Tinaja, 2 6, 1 92, vili.25.62; Papantla, 18 km. E., 2 6, 1 9, ix.9.64; Paso
del Toro, 15 km. W., 1 6, 1 9, ix.8.64, JRZ (NMSU). Poza Rica, 9 m. W.,
26,1 2, vili.27.65, P. J. Spangler (USNM). Puenta Jula, 1 ¢, 2 9, xii.18.48,
H. B. Leech (CAS). Santiago Tuxtla, 10 km. S., 2 6, 2 2, ix.8.64, JRZ
(NMSU).

Laccophilus youngi, new species (Figs. 114-121, 305)

DIAGNOSIS. — The unique combination of a non-irrorated, variegated, or
marmorated elytral pattern, metacoxal file in the male, intermediate prosternal
process, and size makes this one of the most distinctive species in North America.
Only L. pictus and L. gentilis suavis are not irrorated and possess files; but the
first has a black and yellow pattern, and the second is a much smaller species
with a subbasal fascia. In a total of 50 females an epipleural flange was always
present. Other Laccophilus that possess the character that frequently are all
irrorated — L. maculosus shermani, L. fuscipennis, and L. vacaensis chihuahuae.
L. quadrilineatus and L. raitti have variegated or marmorated patterns; but files
are lacking, and the females have rakelike instead of sawlike ovipositors.

DESCRIPTION. — Medium to large (length, 4.6 to 5.4 mm; width, 2.7 to

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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JAMES R. ZIMMERMAN 141

3.0 mm), variegated, light and dark brown species; coxal file present; prosternal
process intermediate; ovipositor sawlike. COLOR. Head: pale brownish-
yellow above and beneath except for reddish-brown occiput; appendages pale
except mandibles. Pronotum: mostly pale brownish-yellow, but with large dark
reddish-brown to nearly black blotches on the anterior margin between the eyes
and continuous with the occipital dark area; posterior margin translucent brown-
ish-red. Elytra: large dark brown or nearly black blotch over most of the disc
on a pale brownish-yellow background; pattern tending to be marmorated with
anterior and posterior extensions from the central blotch; epipleura generally
pale except for the light reddish-brown translucence in posterior half. Tergite
VIII: pale yellowish-brown. Venter: generally light yellowish-brown with
slightly darker reddish-brown tinges at coxal bases and on the hind legs. Geni-
talia: variably reddish-yellow brown. ANATOMY. Méicroreticulation: double
on head, pronotum, and obviously so on elytra. Head: supraclypeal seam di-
verging upward at midline instead of remaining parallel to margin. Pronotum:
epipleural flange nearly always present on females; apical truncation slight in
females; not clearly distinguishable in males. Elytra: WH/PW, 0.68; LP/PW,
0.41. Venter: coxal file prominent in males, composed of about 19 coarse
lines; prosternal process intermediate with apex reaching almost to a line drawn
even with the posterior margins of the mesocoxal cavities and with a sharply
defined crest on apical expanded portion; lobes of postcoxal process rounded,
projecting laterally well beyond the midline; last visible abdominal segment of
male evenly rounded, with numerous prominent rugae and a weakly defined,
nearly symmetrical ventral crest; scattered setigerous punctures and a few thick,
short hairs at ventral lateral margin; female with the apex tending to be triangu-
lar in outline and with a groove or impressed line forming a margin on either
side of the apex; rugae, setigerous punctures and hairs as in male. Legs: pro-
and mesotarsi enlarged in a dorsoventral plane; fifth tarsal segments about one
and one-half to one and two-thirds as long as corresponding fourth; profemoral
setae (7) about half as long as mesofemoral (7 to 9) ones; longer metatibial
bifid spine nearly as long as first two metatarsal segments. Genitalia: oval
plate produced to point; its ventral crest apparent near tip, numerous weak lines
on either side of crest and diverging away from midline; aedeagus widened at
about three-quarters its length, then narrowing to a blunt tip; ovipositor with
about 13 pair of sawlike teeth.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— Laccophilus youngi occurs on the western side of the continent
from Nayarit to Guatemala (Fig. 23). There is also a single male
from Poza Rica, Veracruz. It is found in tropical deciduous forest
at about 2000 to 3000 feet, and I have collected it almost entirely in
deeply cut mountain valley streams with strong current. This prob-
ably accounts for its rarity in collections since most Laccophilus
avoid strong currents. It is also one of the few species that occurs

MEM. AMER. ENT. SOC., 26

142 THE GENUS LACCOPHILUS

where there is a heavy shaded canopy over the stream. It uses grassy
margined and gravelly bottomed, silted streams — as well as those
with solid rock bottoms. They apparently can make use of any kind
of debris to maintain their position in the stream.

MATERIAL EXAMINED

Holotype male, allotype, and four male and four female paratypes with the
following data are in the United States National Museum: 3 miles north of La
Resolana (Casimiro Castillo), x.26.66, A. H. Smith and J. R. Zimmerman.
Other paratypes distributed as follows: three males and nine females, 29 miles
northeast of Colima, Colima (in Jalisco), xii.3.48, H. B. Leech, and two males
and one female, Sierra de Zapotan, xi—.42, Eugenio Paredes, are in the Cali-
fornia Academy of Science, San Francisco, and a male and female, same data,
in the University of Michigan, Museum of Zoology, Ann Arbor; and one male
and one female with the same data as the holotype, Departmento de Ento-
mologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F., Mexico.

MEXICO. — CHIAPAS. Arriaga, 1 2, vii.22.65, P. J. Spangler (USNM).
Cuahtemoc, 1 6, 1 92, viii.30.63, JRZ (NMSU). COLIMA. Trapechi, 1 °,
vil.29.62, JRZ (NMSU). JALISCO. Autlan, 9 m. SW., 4 6, 7 2, x.24.66,
A. H. Smith & JRZ (NMSU). Colima, 29 m. NE. (given as state of Colima,
but this would have to be Jalisco), 3 6, 9 2, xii.3.48, H. B. Leech (CAS).
La Huerta, 5 m. N., 3 2,6m.N., 1 6, 6 2, x.25.66; 22 m. N., 12 6, 14 2,
x.26.66, A. H. Smith & JRZ (NMSU). NAYARIT. Sierra de Zapotan, 3 ¢,
2 2, xi—.42, Eugenio Paredes (CAS). OAXACA. Tapanatepec, 2 ¢, ix.1.63,
JRZ (NMSU). VERACRUZ. Poza Rica, 1 ¢, viii.27.65, P. J. Spangler
(USNM).

GUATEMALA. — Quiroga, 1 92, vili.14.65, P. J. Spangler (USNM).

LACCOPHILUS PICTUS

This polytypic species is composed of three races which have
ranges that extend from the southwestern United States to Guatemala.
Each race has been considered a separate species. L. pictus Castel-
nau was described in 1834 from Mexico, L. insignis Sharp in 1882
from Texas, and L. coccinelloides Régimbart in 1889 from Arizona.
Intergrade populations have been found between pictus and insignis
in Veracruz and between pictus and coccinelloides in Jalisco. Inter-
gradation is restricted to two very small areas.

Although L. pictus males possess metacoxal files, the species ap-
pears to be rather far removed from other North American species
with files. The bright yellow and black elytral pattern contrasts sharply
with the dull irrorated brown or dark brown found in most of the
others. Other differences are the strongly double microreticulation

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MEM. AMER. ENT. SOC., 26

146 THE GENUS LACCOPHILUS

and the long pronotal apex. The relative length of the pronotum
exceeds any other North American Laccophilus.

DESCRIPTION. — Medium to large (length, 4.2 to 5.9 mm; width, 2.5 to
3.4 mm), black and yellow species; metacoxal file present; prosternal process
short; ovipositor sawlike. COLOR. Head: pale brownish-yellow in front; occi-
put, inner eye margin and mandibular margins black; other appendages yellow.
Pronotum: disc to lateral margins pale brownish-yellow, but entire anterior and
posterior margins black filling in the entire “V” at the apex. Elytra: strikingly
marked with yellow and black; epipleura black or dark reddish-brown. Tergite
VIII: pale yellow-brown tinged with red. Venter: entirely pale in shades of
brownish-yellow variably tinged with red. Genitalia: same as venter. ANAT-
OMY. Microreticulation: strongly double on head, pronotum and elytra; indi-
vidual cellules often not visible; pronotum roughened. Head: supraclypeal
seam closely parallel to margin. Pronotum: relatively longer from head to
apex due to the more acute angle formed by the adjacent margins of the pro-
notum and elytra; WH/PW, 0.67; LP/PW, 0.44-0.46. Elytra: epipleural flange
common in insignis, but rare in the other two races; truncation of apices marked.
Venter: coxal file prominent in males and weakly present in females, composed
of about 25 relatively coarse lines, and about 8 to 10 much finer ones; pro-
sternal process with well-defined crest in apical half; lobes of postcoxal proc-
esses rounded and laterally projecting well posterior to midline; last visible
ventral abdominal segments not truncated (discussed under subspecies). Legs:
male pro- and mesotarsi noticeably enlarged in a dorsoventral plane; first three
segments with lateral distal corners produced; palettes large enough to be easily
distinguished at 20 power magnification; fifth tarsal segments on front and mid-
dle legs about one and two-thirds as long as corresponding fourth; mesofemoral
setae (6 to 8) about twice as long as profemoral ones (6 to 7). Genitalia: oval
plate with produced acuminate tip and with prominent ventral crest extending
anteriorly and bending slightly to the left; numerous raised lines on either side
of the crest, but longer and more numerous on the right; right paramere less
triangular than in maculosus (fig. 129) and nearly as long as left; aedeagus
narrowing abruptly near the tip; ovipositor with about 12 sawlike teeth.

Laccophilus pictus pictus Laporte de Castelnau, new status
(Figs: 122-124) 126, 127, 130) 133,13 Seas 09)

Laccophilus pictus Castelnau, 1835, p. 104. Type: unknown, Mexico. Aubé,
1838, p. 441; Horn, 1871, p. 330; Crotch, 1873, p. 400; Sharp, 1882a, p.
290; Sharp, 1882b, p. 11; Horn, 1883, pp. 277, 283; Leng, 1920, p. 77;
Zimmermann, 1920, p. 23; Blackwelder, 1944.

DIAGNOSIS. — The combination of an elytral pattern with bright yellow
spots on a black background, dark epipleura, and male metacoxal file separates
pictus from all other North American species. The principal difference between
p. pictus and p. coccinelloides is that the former has less yellow on the elytra;

JAMES R. ZIMMERMAN 147

L. p. insignis has the yellow spots expanded and fused into zig-zag fascia.
L. horni has a black and yellow elytral pattern, but the epipleura are pale and
the file is absent.

NOMENCLATURAL NOTES. — There has been some confu-
sion in the discussion of the subspecies of pictus; but fortunately there
have been no synonymical complications and no problems related to
other species. The identity of p. pictus itself is not certain, however,
since the type appears lost; and the type locality was given only as
Mexico, and all three races occur there. I have not designated a neo-
type because I have not, as yet, visited the European museums where
the type might be deposited. The type locality is being restricted,
however, to El Cameron, Oaxaca.

VARIATION. — The races of pictus are probably the most strik-
ingly patterned North American Laccophilus. The strongly contrast-
ing and well-defined areas of yellow and black on the elytra permit
easy and concise comparisons between individuals and populations.
There is some individual variation in the size of the discrete spots
and the degree to which other parts of the pattern are interconnected.
There is a tendency for the second and fourth spots along the sutural
midline to be smaller or absent in p. pictus. There is no apparent
sexual difference in the elytral pattern.

Females are larger than males (Table 10, Fig. 14). The differ-
ence in mean size is more than 0.25 mm. The largest male from a
sample of 317 specimens is 5.08 mm long, and the largest female in
a sample of 371 specimens is 5.24 mm long. Populations from
Jalisco to Honduras are remarkably similar in overall length. Plots
of the frequency distributions for length result in smooth curves for
males and females.

A sample taken about ten miles inland from Autlan, Jalisco,
showed all degrees of intermediacy between pictus and coccinelloides.
But at Autlan only the pictus elytral pattern and aedeagus are pres-
ent in the population. Size, however, approaches the values for coc-
cinelloides, indicating there is some introgression that is not apparent
from analysis of elytral pattern or male genitalia. The average sizes
for pictus is 4.60 mm for males and 4.87 for females; but the means
for the Autlan sample is 4.70 for males and 5.01 for females, which
is closer to coccinelloides and is larger than any other samples of pic-

MEM. AMER. ENT. SOC., 26

148 THE GENUS LACCOPHILUS

tus except for those that might be influenced by insignis in Veracruz.

The PW/EL values which range from 0.590 to 0.617 are as high
as those for L. maculosus. The means for males are higher in every
sample, indicating a slight difference in body proportions. In both
sexes, however, the length is 1.69 times the width, which means the
shape of the pronotum may be relatively shorter in males. Another
answer is that the greater curved surface of the larger females is cre-
ating a parallax error in measurement of the elytral length.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— L. p. pictus ranges from Jalisco to central Veracruz and south to
Honduras. It intergrades with coccinelloides between Autlan and
Guadalajara in Jalisco and with insignis near Papantla, Veracruz
(Fig. 13). It is most commonly found at elevations between 1000
and 4000 feet in arid tropical scrub in small mountain streams and
pools that have granitic gravel bottoms. The bright yellow and black
elytral pattern may be some protection in the clear streams that have
irregular sized grains on the bottom. It is rare on the coastal plain.
It will sometimes be abundant in grassy, mud-bottomed temporary
ponds or puddles during the rainy season.

MATERIAL EXAMINED

MEXICO. — CHIAPAS. Arriaga, 1 ¢, 1 2, vii.22.65, P. J. Spangler (USNM).
Chiapa de Corzo, 1 4, ix.1.63; Cintalapa, 5 m. W., 1 6, 4 2, ix.1.63; Comitan,
1 3, vii.30.63; Cuahtemoc, 4 m. W., 1 4, viii.30.63; Ixtapa, 3 ¢, 2 2, vili.31.63;
Ocozucuatla, 6 m. W., 1 6, 1 2, ix.1.63; Tuxtla Gutierrez, 5 3, 3 2, ix.1.63,
JRZ (NMSU). Las Cruces, 5 m. S., 21 4, 28 9, viii.23.65, P. J. Spangler
(USNM). COLIMA. Colima, 7 m. NE., 24 6, 28 2, xii—.48, H. B. Leech
(CAS). Manzanillo, 5 m. S., 1 ¢, 1 2, vii.29.62; Tecoman, 5 m. N., 1 @,
vii.29.62; Trapechi, 9 6, 13 2, vii.29-30.62, JRZ (NMSU). GUERRERO.
Mexcala, 1 9, i.8.56, E. C. Bay (CNL). Zumpango, 12 m. N., 12, 1.8.56,
J. C. Schaffner (ISU). JALISCO. Autlan, 62 6, 42 2, x.26.66; 9 m. SW.,
25 6,39 9, x.24.66; Barra de Navidad, 2 6,5 2, x.25.66; La Huerta, 6m. N.,
25 6, 39 9, x.25.66;5 m.N., 2 6,4 2, x.25.66; 6 m. S., 39 6, 40 9, x.24.66;
9m.S.,3 6,3 2, x.25.66, A. H. Smith & IRZ (NMSU). Tecalitlan, 10 m. S.,
4 9, iii.27.64, JIRZ (NMSU). MEXICO. Tonatico, 13 ¢, 8 9, vili.29.62,
JRZ (NMSU). MICHOACAN. Apatzingan, 1200 feet, 1 2, viii.11.41, H.
Hoogstraal (CAS). Uruapan, 2 4, iii.26.64; Tzitzio, 3 m.S., 2 6, 1 2, vii.22.63;
1 6, vii.27.62, JRZ (NMSU). OAXACA. El Cameron, 27 ¢, 34 9, viiil.27.63;
25 km. N., 16 6, 21 92, ix.6.64; El Coyula, 16 km. S., 7 6, 6 2, ix.6.64; Hua-
juapan, 2 km. S., 5 4, 5 9, ix.4.64; 17 km. NE., 1 6, ix.3.64; Juchitan, 20
km. N., 1 2, ix.7.64; Matias Romero, 5 m. S., 1 9, 1x.7.64; Oaxaca, 2 6, 1 2,
vili.25.63; Tehuantepec, 1 ¢, 2 2, ix.6.64; Texquisitlan (Rd. 190), 9 4, 7 9,

JAMES R. ZIMMERMAN 149

LACCOPHILUS P INSIGNIS 4

LACCOPHILUS P.
COCCINELLOIDES o

Che

Figure 13. Distribution of Laccophilus pictus.

MEM. AMER. ENT. SOC., 26

150 THE GENUS LACCOPHILUS

ix.6.64; Tlacolula, 1 ¢, ix.6.64, JRZ (NMSU). Amatitlan, 3 6,9 9, xii.10.48,
H. B. Leech (CAS). PUEBLA. Puebla, 2 6, 1 9, vi.26.57, D. R. Lauck
(USNM). Tehuitzingo, 15 ¢, 17 9, viii.24.63, JRZ (NMSU). VERACRUZ.
Cuitlahuac, 9 ¢, 11 92, viii.25.62; Huatusco, 25 km. S., 7 6, 1 2, ix.9.64; La
Tinaja, 4 8,2 9, vili.25.62, JRZ (NMSU). Paso del Macho, 25 km. NE., Cor-
doba, 1 6 (from B.C.A.), Hoege, donated by F. DuGodman, 1907 (AMNH);
2 ¢ (same locality data), (ANSP). Paso de Ovejas, 2 $, 6 Q, viii.27.62, JRZ
(NMSU).

EL SALVADOR. — Usulutan, 5 m. E., 1 2, vii.31.65, P. J. Spangler (USNM).
GUATEMALA. — Aldea Jesus Maria, 4 6, 8 9, viii.15.65; El Progreso, 5 m.
S., 16,5 @, viii.11.65; Pijije, 1 6, vii.8.65, P. J. Spangler (USNM).
HONDURAS. — Choluteca, 10 m. W., 1 @, vii.29.65; Jicaro Galan Jct., 2 2,
vil.29.65; Pespire, 12 6, 15 2, vii.29.65; Sabarra Grande, 10 m. N., 1 ¢,
vii.29.65; San Marcus Colon, 39 6, 29 @, vii.25.65, P. J. Spangler (USNM).

NICARAGUA. — Esteli, 9 m. N., 4 ¢, 4 9, vii.10.65; San Benito, 13 m. N.,
1 Q, vii.11.65; Somoto, 6 6,5 2, vii.10.65; 4 m. W., 6 6,5 9, vii.10.65, P. J.
Spangler (USNM). Musawas, 1 ¢, xi.1.55, B. Malkin (CAS).

Laccophilus pictus coccinelloides Régimbart, new status
(Figs. 130, 132, 136, 300)

Laccophilus coccinelloides Régimbart, 1889, p. 112. Type: Leyden Museum,
Arizona. Leng, 1920, p. 77; Zimmermann, 1920, p. 17.
Laccophilus pictus Leech, 1948, p. 400.

DIAGNOSIS. — The same combination of characters for p. pictus applies
for p. coccinelloides; the principal differences are in the structure of the male
aedeagi and in the presence of four larger yellow spots near the elytral midline
of the latter.

NOMENCLATURAL NOTES. — The identity of Régimbart’s
coccinelloides (unlike Castelnau’s pictus) is certain since Arizona
was given as the type locality. The race from southern Mexico and
Central America was retained as typical pictus. Also, there can be
no confusion of insignis and coccinelloides since the former occurs
no farther west than Texas. Horn (1883) and Leech (1948b). used
pictus for the populations in Baja California, but coccinelloides is the
race that occurs there.

VARIATION. — There is no marked sexual difference in pattern
and no recognizable trend in the pattern variation, but females aver-
age more than a 0.25 mm larger than males (Table 11, fig. 14).
There is also no clear geographic size variation. The two largest
groups are females from the cape region of Baja California and from
the Davis Mountains of Texas (Table 10), and the smallest males

JAMES R. ZIMMERMAN 151

|

insignis

Ye

pictus

U M B E R §S

N

coccinelloides

5.4 5.6 5.8

TOTAL LENGTH (MM)

Figure 14. Histograms of length in Laccophilus pictus. Males are shown
crosshatched; females, stippled.

come from northeastern Sonora and from Nayarit. Plots of the fre-
quency distribution of total length suggest, however, that there is some
unanalyzed heterogeneity in the samples. The peak occurs about 4.8
mm, but the mean is 4.68 mm; and the slope is irregular, with the sug-
gestion of another peak at less than 4.6 mm. This curve contrasts
with the smooth one obtained for pictus. To some extent, the samples
from Baja California and the Davis Mountains of Texas are isolated

MEM. AMER. ENT. SOC., 26

SZ THE GENUS LACCOPHILUS

from the rest of the subspecies; and it might be that, with sufficient
material, they would prove to be statistically different populations,
contributing to the irregularity of the frequency distribution curve.

L. p. coccinelloides appears to be slightly larger than p. pictus
and smaller than p. insignis. The sample from near Union de Tula
has the size of coccinelloides, but the elytral pattern and aedeagus
show all degrees of intermediacy. The WP/EL ratio is the same for
both races.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This race occurs from central Arizona to central Jalisco and to the
extreme southern end of Baja California (fig. 13). It is also common
in the extreme southwestern county (Hidalgo) of New Mexico and in
west Texas in the Guadalupe, Davis, and Chisos Mountains. It should
occur in southern California. While it is more characteristic of the
west slope of the Sierra Madre Occidental, it is easy to find on the
Plateau in Chihuahua and Durango. It and pictus occur mostly at
1000 to 4000 feet, but can be found below and above those limits.
Its usual habitat is in pools of gravelly bottomed mountain streams in
the pine-oak woodland vegetation zone.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Douglas,
1 $, 1.15.33: 1 2, iv.2.33, W. W. Jones (BERK). Huachuca Mins., 21,
v.7-8.53, A. H. Dietrich (CNL). Portal, 5 m. W. SWRS, 2 9, x.1.55, M.
Cazier; 2 2, v.4.56, M. Statham (AMNH); Portal, 1 2, vi.4.59, L. A. Stange
(DAV). Navajo County. Near Kayenta, Tsegi Can., 3 m. N., Marsh Pass,
1 6, ix.13.50, J. Figg-Hoblyn (CAS). Pima County. Baboquivari Mtns.,
Brown Can., 1, viii.8.53, D. Butler (ARI); Baboquivari Mtns., 3, —.—.23; 6,
x. 1-15.24, O. C. Poling (CNL); 2, viii.18.32, D. K. Duncan (CNL). Organ P.
Cactus N.M., Ajo Mtns., Alamo Can., 1 6, xii.14.39, G. G. Habison (CAS).
Santa Catalina Mtns., Bear Canyon, 9 6, 15 9, i.2.38, Van Dyke (CAS); 1 4,
2 9, 11.15.64, J. W. Green (CAS); 3, xi.10.36, L. P. Wehrle (ARI); 1 6,3 2,
ii.1.38, J. W. Tilden (FM); Molino Basin, 1 6, 1 2, vi.9.54, R. S. Beal (CAS);
16, 1 9, vi.9.54, R. S. Beal (BERK); 1 6, vi.11.58, C. D. & F. MacNeil
(CAS); Lowell Rngr. Sta., 7, vii.6-20.16 (USNM); 11, vi.18.—, Hubbard &
Schwarz (USNM); Sabino Canyon, 1, x.26.15, Dodge (USNM); 1 2, J. W.
Tilden (FM); Santa Catalina Mtns., 1 2, v.8.33, O. Bryant (CAS); 2, x.6.42,
E. Brainard (ARI). Tucson, 1 4, iv.10.38 (FM); 1 4, 5 8, iii.13.46, J. W.
Green (DAV); 5 6, 11 92, i.27.35, O. Bryant (CAS); 3 9, ix.-x.—.28; 2 2,
ix.—.28, F. H. Parker (CAS); 1 9, x —.28, F. H. Parker (BERK); 2, iii.13.36,
L. P. Wehrle (CNL); Sabino Canyon, 6, iii.13.36, J. G. Needham (CNL);
1, 11.21.53, A. & H. Dietrich (CNL); Tucson, 15 m. NE., 2 6, 3 @, x.5.63,

JAMES R. ZIMMERMAN 153

K. L. McWilliams (NMSU). Pinal County. Oracle, 14 m. E., 3 6, 2 2,
vii.27.24, J. O. Martin (CAS). Santa Cruz County. Atasco Mtn., 1, vii.27.54,
D. Butler (ARI); Coronado N.F., Pena Blanca L., 4 4, ix.6.61, JIRZ (NMSU);:
Nogales, 1 6, 2 2, iv—.97, Koebele; 3 ¢, 1 2, ix.3.06, 2 6, viii.31.06, F. W.
Nunenmacher, 1 ¢, vili.19.06 (CAS); 1 46, ix.3.06 (FM); 1 °, ix.3.06, F. W.
Nunenmacher (CAS); Pena Blanca, 1 Q, vii.1.61, 2 4, vii.11.61; 1 9, vii.31.61;
1 2, vili.4.61; 1 9, viii.9.61; 1 6, 1 9, viii.11.61; 2 6, 9 2, viii.12.61; 4 9,
ix.6.61; 1 6, 3 2, vi.28.62; 1 9, vii.12.62; 2 @, vii.14.62; 1 2, vii.24.62; 2 9,
vili.8.62; 1 6, vii.12.63; 1 2, vii.23.64, R. H. Arnett & E. Van Tassel (CUA).
Patagonia, 3 m. S., Sonoita Ck., 2 6, 3 2, ix.6.61, JRZ (NMSU). Sycamore
Cn., Tumacacori Mt., 1,6, 7 2, vii.27.65, H. B. Leech (CAS). Ruby, 2,
xi.20.55, F. Werner & G. Butler (ARI). Santa Rita Mtns., Box Can., 2 é,
vill.29.52, B. Malkin & V. Thatcher; Santa Rita Mtns., 1 2, x.21.34, O. Bryant
(CAS); 1, xi.18.—, Hubbard & Schwarz (USNM). Tumacacori, Sycamore
Can., 1, vi.3.53, A. & H. Dietrich (CNL). Yavapai County. Castle Hot
Springs, 2 ¢, 2 2, iv.5.42 (AMNH). Bloody Basin (about 25 m. S. of Verde
Hot Springs), 2 6,7 92, vi.8.47, F. H. Parker (CAS). Prescott, 2 2, vili—m.—
(AMNH). NEW MEXICO. Hidalgo County. Animas Mtns., Double Adobe
Rnch., 1 2, viii.15.52, H. B. Leech (CAS). Dona Ana County. Las Cruces,
Rio Grande R., 1 2 (NMSU). TEXAS. Brewster County. Alpine, 20 m. S.,
1 $, v.12.17, J. O. Morton (CAS). Rio Grande(?), 5, vi.13-17.08, Mitchell
& Cushman (USNM). Big Bend Ntl. Pk., Chisos Mtns., 1 2, vii.3.42, H. A.
Scullen (ORES). Culberson County. Nickel Ck. Sta., 2.5 m. E., 1 6, ix.9.52,
B. Malkin (CAS). Davis County. Davis Mtns., Elbow Canyon Ck., 6 ¢,
8 2, x.27.61, JRZ (NMSU). Limpia Canyon, 1, iv.19.53, B. Adelson & M.
Washbauer (BERK); 17 6, 34 92, ix.4-5.52, B. Malkin (FM). Madera Can-
yon Ck., 1 6, viii.3.61; 1 2, x.27.61; Davis Mtns., tank, 1 6, x.27.61, JRZ
(NMSU). Ft. Davis, 2 2, iv.19.53, Adelson & Washbauer (USNM); 1 2,
vili.10.16 (MCHS); 1 m. N., 4, vii.16.41, B. E. White (CAS). Kent, 13 m. S.,
1 6, 1 &, 1x.8.52, B. Malkin (FM). Old Ft. Canon, 1, 1ii.8.36, J. G. Need-
ham (CNL).

MEXICO. — BAJA CALIFORNIA. Aqua Caliente (Cape region), 9 6, 14 9,
iv.22.47, I. LaRivers (BERK). Cabo San Lucas, 7.7 m. NE., 1 ¢, 1 2, 1.6.59,
H. B. Leech; El Triunfo, 7 ¢, 9 Q, viii..13.38, Michelbacher & Ross; Miraflores,
3 m. NW., 1 2, 1.19.59, H. B. Leech; La Parras, 1 6, x.23.—, W. M. Mann;
La Suerte, 1 ¢, vi.4.63, R. K. Benjamin; La Paz, 12.4 m. E., 2 6, 2 9,
xii.23.58, H. B. Leech; 18.5 m. S., 1 6, 1 2, xii.19.58, H. B. Leech (CAS).
Padarone, 7 ¢, 6 2, v.11.47, Ira LaRivers; Todos Santos, 2 ¢, 1 2, iv.10.47,
I. LaRivers; 1 2, iv.10.47, Ira LaRivers; San Bartolo, 6 6, 1 2, v.1.47, I.
LaRivers (BERK). 1m. SE., 1 9, i.20.59, H. B. Leech; San Antonio, 1 é,
1 9, vii.12-17.19, J. R. Slevin; 1 2, vii.12.19, G. F. Ferris; San Jose del Cabo,
1 ¢, 12 (CAS). San Luis Gonzaga,12 m. E., 1 ¢, 2 9, v.22.47 (BERK).
San Felipe, 2 ¢, G. W. Beyer (CAS); 8 3, 14%, G. W. Beyer (AMNH).
Santa Rosa, 1 4, 2 9, viii.-ix—.01, G. Beyer (FM). Sierra Juarez, Guada-

MEM. AMER. ENT. SOC., 26

154 THE GENUS LACCOPHILUS

lupe Cn., 1 6, 1 9, vii.6.55, R. Orr & C. Tese (CAS). CHIHUAHUA. Chi-
huahua, 43 m. N., 2 6, 1 2, xii.8.62, JRZ (NMSU). Chihuahua, 12 m. N.,
1 2, viii.23.60, Arnaud, Ross, Rentz (CAS). Parral, 2 m.S.,3 4,1 2, vii.25.62,
JRZ (NMSU). DURANGO. Abasolo, Rio Nazas, 1 ¢, x.22.66, A. H. Smith
& JRZ; La Zarca, 15 m. N., 1 ¢, vii.25.62; San Juan del Rio, 10 m. N., 1 4,
1 2, xii.9.62, JIRZ (NMSU). JALISCO. Barranca Oblato, 8 m. NE. Guada-
lajara, 53 6, 34 9, A. H. Smith & JRZ; Guadalajara, 20 m. E., 1 9, iii.25.64,
JRZ (NMSU). Tala, 4 6,5 2, iii.25.64, JRZ; Tecolotlan, 1 ¢, x.24.66, A. H.
Smith & JRZ (NMSU). Tlaquepaque, 2 2, vii—.53, N. L. H. Krauss (CAS).
NAYARIT. Tepic, 25 km. S., 3 ¢, 3 9, ix.24.53, B. Malkin; 24 m. SE., 1 ¢,
1 2, vili.16.60, Arnaud, Ross, Rentz; Ixtlan del Rio, 9 ¢, 14 9, ix.22.53, B.
Malkin; San Blas, 1 9, ix.17-21.53, B. Malkin (CAS). SINALOA. Elota,
1m. S., 1 6, 4 9, viii.1.62; Concordia, 8 m. E., Rd. 15, 1 9, xii.12.62; Ma-
zatlan, 7 m. S., Rd. 15, 1 9, xii.11.62, JRZ (NMSU); 72 m. N., 1 6, i.—.62,
Breedlove & Copp (CAS). SONORA. Alamos, 1 2, vii.30.40, R. P. Allen
(CAS); 1 2, xi.1.60, R. L. Westcott (LACM); Aribabi (E. of Moctezuma),
30 6, 65 9, xii.15.62, JRZ (NMSU). 7m. SE., 7 ¢, 6 9, viii.12.60, Arnaud,
Ross, Rentz (CAS). Esqueda, 20 m. S., 2 2, vi.26.56, F. N. Young (FM);
Guaymas, 1 6, ix.29.63, W. M. Mann (CAS). Nogales, 42 km. S., Rancho
Atascosa, 1 6, 1 @, ix.21.52, B. Malkin (FM).

Laccophilus pictus insignis Sharp, new status
(Figs. 125, 128, 131, 134, 138, 302)

Laccophilus insignis Sharp, 1882a, p. 290. Type: male (?), British Museum
(Natural History), Texas. Horn, 1883, p. 277; Zimmermann, 1910, p. 20;
Leng, 1920, p. 77; Blackwelder, 1944, p. 74; Leech, 1948, p. 401.

DIAGNOSIS. — The black and yellow elytral pattern with a yellow zig-zag,
subbasal band separates insignis from all other North American Laccophilus.

NOMENCLATURAL NOTES. — There has been some confu-
sion of insignis with other races of pictus, but not with other species.

VARIATION. — Unlike pictus and coccinelloides, insignis fe-
males are smaller than males (Table 10, fig. 14). In all five locality
samples the lengths of males exceeded that of females. The differ-
ence is about 0.07 mm. The WP/EL ratio appears to be slightly
higher for males. The elytral pattern appears to be fairly uniform
throughout the range, and there is no sexual difference. Intergrades
between pictus and insignis from near Papantla, Veracruz, showed all
degrees of pattern variation between typical pictus and insignis, but
were close to the mean of insignis in size.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L. p. insignis ranges from west Texas to central Veracruz mainly
on the eastern slope of the Sierra Madre Oriental (fig. 13). It is one

JAMES R. ZIMMERMAN 155

of the most common Laccophilus in the hill country of central Texas
(Blanco County, for example). It is usually taken between 500 to
3000 feet in stream pools. It prefers granitic gravelly bottomed pools
which frequently occur in oak woodland in the northern part of the
range and in tropical deciduous or tropical evergreen forest in the
southern part.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — KANSAS. Sedgwick County. Derby,
1 2, vii.15.58, JRZ (NMSU). OKLAHOMA. Greer County. Mangum, 1,
ix.9.— (USNM). TEXAS. Blanco County. Cypress Mill(s), 1, iv.2.—; 2,

ix.10.88; Round Mountain, 1, —.—.26, H. S. Bolier; 1 (USNM); 2 6, 1 2
(ANSP); 5 6, 1 2 (MCZ). Shovel Mtn., vi.—.26; 1, —.—.29; 2 (USNM);
1 (CNL); 18 6, 27 2, ix——, F. G. Schaupp (AMNH); 3 6 (MCZ). Bur-

net County. Marble Falls, 1, iii.14.59, E. O. Morrison (TAM). County, 2
(USNM). Colorado County. Columbus, 3 6, 42 (CARR). Coryell County.
Fort Hood, 1, vi.5.55; 1, vii.19.55; 3, vii.30.55, Matthews (CNL). Falls
County. Reagan (Wells?), 4, iii.6.36,J.G. Needham (CNL). Gillespie County.
Fredericksburg, 5, vi.22.55, W. W. Boyle (CNL). Hays County. Dripping
Springs, 3, viii.9.42, W. S. Ross & E. S. Ross (CAS). Kendall County. Com-
fort, 1 6 (CARR). Kerr County. Kerrville, 3, x.8.05; 5, vi.19.07, F. C. Pratt
(USNM); 2, vi.21.42, E. S. Ross (CAS). Llano County. Llano, 2 6, 4 2,
iv.25.63, George Child (NMSU). Sutton County. Sonora, Dry Devil’s River,
5, xi.5.49, O. Bryant (CAS). Travis County. Austin, 3, vi.28.33; 5 (USNM).
Lake Austin, 1 2, iv.3.53, J. E. Gillespie (AMNH). Uvalde County. Sabinal, 1,
vi.19.10, F. C. Pratt (USNM). Uvalde, 3, v.21.33 (TAM). Val Verde
County. Del Rio, 1 6, 1 9, vil.23.24 (MCZ). Zapata County. 1, ii.26.36,
J. G. Needham (CNL). Dubious record. El Paso County. El Paso, 4 6,5 2
(AMNH#).

MEXICO. — COAHUILA. Ramos Arizpe, 1 6, 2 9, vii.7.63, JRZ (NMSU).
NUEVO LEON. Allende, 1 ¢, vii.8.63, JRZ (NMSU). Sabinas Hidalgo,
1 4, xii.16.40, F. N. Young (AMNH). Sta. Catarina, Huasteca Can., 1 é,
vii.7.63, JRZ (NMSU). SAN LUIS POTOSI. Agua Zarca (N. Platinito),
7 3, 3 Q, 11.25.63; Antiguo Morelos, 3 9, iii.23.63, JRZ (NMSU). Cuidad
del Maiz, 15 m. E., 1 92, xi.19.48, H. B. Leech. El Salto, 5 9, iii.25.63, Guay-
muchil (N. Naranjo), 3 6, 11 9, iii.25.63; Jitalpa, 9 ¢, 5 2, ili.23.63, JRZ
(NMSU). Palitla, 5 m. N. of Tamazunchale, 3 ¢, 8 92, xii.22.48, H. B. Leech.
Valles, 29 m. N., 1 6, viii.19.54, F. N. Young (AMNH). TAMAULIPAS.
Antiguo Morelos, 3 m. N., 1 6, 5 9, iii.26.63; 6 2, El Limon, 1 6, 2 2,
11.24.63; Llera, 1 6,5 92, iii.23.63; Mante, 1 2, iii.23.63; Nuevo Morelos, 5 4,
9 2, iii.25.63; Ocampo, 6 6, 8 2, ili.24.63, JRZ (NMSU). San Jose, 2 4,
iv.i—.10 (USNM). Victoria, 3 é.2 2, xii.10.—, F. C. Bishop (USNM).

MEM. AMER. ENT. SOC., 26

156 THE GENUS LACCOPHILUS

L. p. pictus X L. p. insignis
VERACRUZ. Papantla, 18 km. E., 37 6, 31 2, ix.9.64, JRZ (NMSU).

LACCOPHILUS VACAENSIS

Laccophilus vacaensis Young is the most recently described Lac-
cophilus in North America. It was described from a single locality
on Key Vaca, Florida (1953). Specimens from other localities have
long been present in the material from several museums; but due to
its similar appearance in color, pattern, size, and shape to L. proxi-
mus, it was not recognized as distinct. It has a wide distribution and
may be present in every one of the states along the southern tier of the
United States, as well as occurring over much of Mexico and all the
way to Costa Rica. It appears related to L. tarsalis Sharp, a South
American species. There are three geographical races in North Amer-
ica. The most widely distributed is v. vacaensis, which has a perplex-
ing distribution and may prove to be composed of more than one race
when more is known about it. L. v. chihuahuae occurs from south-
central Texas to southeastern Arizona. L. v. thermophilus is distrib-
uted around the Gulf of California. As yet, no intergrades are known
between the races; but so few specimens have been taken from the
possible intergrade areas that it cannot be concluded they do not inter-
grade. Apparently, all three races occur in southeastern Arizona;
and it is possible that there is a circular overlap without intergrada-
tion in that region. This species appears to be most closely related to
L. spangleri with which it must be considered sympatric in southern
Mexico and Central America; but the collections are, unfortunately,
so scattered that this is also an open question.

DESCRIPTION. — Medium (length, 4.0 to 5.3 mm; width, 2.2 to 3.0 mm),
brown, irrorated species; metacoxal file absent; prosternal process short; fork-
like ovipositor. COLOR. Head: generaily pale brownish-yellow, but often
with reddish tinge especially on the occiput between the eyes; appendages also
yellow. Pronotum: generally the same as the head, but with darker brown
areas of varying intensity at the anterior margin between the eyes and at the
apex. Elytra: irrorated pattern variable and discussed under subspecies; epi-
pleura pale anteriorly, but may darken posteriorly. Tergite VIII: pale yellowish-
brown. Venter: variable from light brownish-yellow to light brown and light
reddish-brown; metacoxal plates and hind legs usually darker than rest of venter.
Genitalia: variable from light reddish-yellow brown to dark reddish-brown.
ANATOMY. Microreticulation: faintly double on the vertex of the head but
most cellules irregularly elongate; pronotum with some cellules double, but most
single and irregularly elongate; single on elytral disc and more deeply impressed

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MEM. AMER. ENT. SOC., 26

158 THE GENUS LACCOPHILUS

in females (pebbled). Head: supraclypeal seam arching slightly upward away
from the margin. Pronotum: WH/PW, 0.66 to 0.67; LP/PW, 0.39 to 0.40.
Elytra: slightly truncated; crease lying just above the female epipleuron and
extending in a straight line posteriorly to just beyond half the length of the
elytra; epipleura with prominent flanges in all females of chihuahuae and ther-
mophilus but only rarely in v. vacaensis. Venter: prosternal process with well-
defined median crest; postcoxal processes laterally projecting posterior to the
midline; last abdominal segment of female not truncated, but subtriangular with
a rounded ventral median crest and a marginate groove on either side and with
scattered setigerous punctures; male last segment not clearly truncated, but ap-
pearing so in some specimens due to flexing on either side of the asymmetrical
median crest; slightly produced at apex, with scattered setigerous punctures;
hind margin of male fifth visible segment with creases or protuberances on
either side of midline, the left more prominent than the right. Legs: male pro-
and mesotarsi swollen laterally as well as enlarged in a dorsoventral plane; fifth
tarsal segments on both pair of front legs more than twice as long as corre-
sponding fourth; palettes easily visible at 20 power magnification; profemoral
marginal setae (10 on male; 7 to 8 on female). Genitalia: oval plate with pro-
duced acuminate tip with ventral crest extending anteriorly with slight curve to
the left nearly to front of plate; prominent raised lines on either side; aedeagus
bent near base and subapically, relatively straight between; ovipositor with
about nine pair of spaced, rakelike teeth.

Laccophilus vacaensis vacaensis Young, new status
(Figs. 139-148, 150, 153, 313, 314)

Laccophilus vacaensis Young, 1953a, pp. 31-34. Holotype: University of Michi-
gan Museum of Zoology, male; near Marathon, Vaca Key, Florida, viii.22.49,
J. S. Haeger and F. N. Young; Young, 1954, p. 46; Young, 1963, p. 5.

DIAGNOSIS. — L. v. vacaensis has an irrorated pattern and color similar
to a dozen other species of North American Laccophilus, but possesses struc-
tural characters in both males and females that permit comparatively easy iden-
tification. The males have the fifth segment of the two front pair of tarsi more
than two and a quarter times as long as the corresponding fourth. Other spe-
cies do not exceed twice the length. Females have a crease, immediately above
the epipleuron, that extends nearly two-thirds the length of the elytron. The
presence of this crease seems to be directly related to the longer tarsal segment
of the males. The distance from the ventral margin of the epipleuron to the
crease is about the same as the length of the fifth tarsal segment, so this is prob-
ably an adaptive feature to assist the male to grasp the female for copulation.

L. v. vacaensis is most likely to be confused with the sympatric species, L.
proximus and L. spangleri. In the first case, the lack of a file in the male
and the presence of a forklike, instead of sawlike, ovipositor in the female will
assist — along with the characters given above. L. spangleri can be best sepa-
rated by the male shorter fifth tarsal segment or by the weaker crease on the
female elytra and by the male aedeagus.

JAMES R. ZIMMERMAN 159

VARIATION. — The degree of pigmentation of the pronotum
and of the elytral pattern varies considerably in this race. Specimens
from Yucatan and Central America are much darker than those from
Texas and the east coast of Mexico, with the individual dots of the
pattern tending to form more continuous chains of five or six dots.
A few specimens tend to be dark enough to suggest suffusion of pig-
ment rather than irroration. Central American specimens have a
prominent darkened area on the anterior margin of the pronotum
between the eyes, while those from farther north may have only a faint
suggestion of reddish-brown or brown color.

Males appear to average 0.05 to 0.10 mm larger than females, but
the largest specimen examined was a female (fig. 16). There seems to
be a geographical difference in size, since the specimens from Guate-
mala and Costa Rica (especially the latter) are almost 0.1 mm larger
than those from south Texas and almost 0.25 mm larger than those
from Progreso, Yucatan. There are too few samples to decide whether
the difference in size is racial, clinal, or local.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— The range of this race is a wide one and a perplexing one. It was
first described from a single locality from the Florida Keys and is now
known to occur on several Caribbean Islands as well. It should occur
clear around the Gulf of Mexico, but it has not been reported from
Alabama or Mississippi. It is found in lowlands of Central America
(fig. 15). This part of its distribution is entirely logical, if one as-
sumes that it occurs in temporary pools and ponds that are often sub-
ject to temperatures above 40° C.; but the rest of the distribution is
more difficult to explain; making it appear that another unrecognized
race is involved. It has been collected from several localities at 5000
feet or above in Morelos, Jalisco, Sonora, and Arizona. It would
thus be expected that these populations should be closer to v. chihua-
huae, which more typically occurs at the higher elevations; but they
are not.

Young (1953a) collected vacaensis in a small freshwater pool in
which the principal vegetation was Chara. The pool was drying up,
the surface was unshaded, and the noon temperature of the surface
water may have exceeded 40° C. I have collected a large number of
specimens from what may have been a brackish temporary pool about
a mile inland from an extensive mangrove swamp near Progreso,

MEM. AMER. ENT. SOC., 26

160 THE GENUS LACCOPHILUS

Yucatan. The water was very dark, only a few inches deep, and filled
with debris, but had some emergent, grassy vegetation. Interestingly
enough, vacaensis was taken in association with L. quadrilineatus
mayae and Young (1963) also described vacaensis as being in associa-
tion with L. inagua Young in the Bahamas. L. inagua most resembles
q. mayae among North American and West Indian Laccophilus.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Santa Cruz County. Pena
Blanca, Pajarito Mts., 2 ¢, 1 2, vii.19.62, Arnett and Van Tassel (CUA).
LOUISIANA. Cameron County. Cameron, 1 @, vii.10-14.05, Van Dyke?
(CAS). TEXAS. Bastrop County. 1 2, vii.13.37 (TAM). Cameron County.
Brownsville, 3, vii—.—; 16, ——.— (USNM); 1 4, ii.24.96, H. F. Wickham
(AMNH); 1 2, x.—.42, E. S. Ross (CAS); 1 6, iim—, Wickham (ANSP);
1 6, Wickham (MCZ). Port Isabel, 1 6, 7 2, x.20.49, O. Bryant (CAS).
Hidalgo County. Edinburg, 1, ii.24.36, J. G. Needham (CNL). Kleberg
County. Kingsville, 4, —.—.—, C. T. Reed (CNL); 1 2, x.20.61, B. Mc-
Daniels; 1 ¢, xi.5.59 (USNM). Victoria County. Victoria, 3, ix.18.14, J. D.
Mitchell (USNM); 1, ix.22.14 (USNM).

BRITISH WEST INDIES.— BAHAMAS. Great Inagua Is., Matthewtown,
8 6,7 Q, 1.31.53, L. Giovannoli & E. B. Hayden; New Providence Is., Nassau,
1 92, 1v.5.53, E. B. Hayden (AMNH).

CUBA. — Santiago, Vista Alegra, 1, vi.16.42, C. T. Randall (USNM).
HAITI. — Attelye, 1 6, 1 2, x.22.25, W. A. Hoffman (USNM).

COSTA RICA. — Bagaces, 2 6, 2 9, vii.12.57, D. R. Lauck; La Cruz, 16 m.
S., 100 +, vii.25.63; Liberia, 12 m. SW., 1 6, vii.25.65; 5 m. SW., 22 6, 31 2,
vii.24.65, P. J. Spangler (USNM).

GUATEMALA. — Aldea Jesus Maria, 6 3, 6 2, viii.15.65, P. J. Spangler
(USNM).

MEXICO. — CAMPECHE. Champoton, 11 m. SW., 1 6, ix.27.63, JRZ
(NMSU). JALISCO. Guadalajara, 11 m.S.,2 6, 1 2, vii.30.62, JRZ (NMSU).
Jiquilpan, 15-20 m. W., 1 6, xi.30.48, E. S. Ross (CAS). Near Cuautla, 1 2,
3 Q, vill.28.62; Puente de Ixtla, 1 2, viii.29.62, JRZ (NMSU). SINALOA.
Mazatlan, 2 4, viii.1.62; 1 2, xii.11.62, JRZ (NMSU). SONORA. . Aribabi
(E. of Moctezuma), 2 6, xii.15.62, JRZ (NMSU). Naco, 1 9, viii.15.49,
G. M. Bradt (AMNH). TAMAULIPAS. Llera, 20 m. S., 1 4, iii.24.63, IRZ
(NMSU). Matamoros, 15 m. S., 1 ¢, vi.10.60, F. N. Young (UMMZ).
YUCATAN. Progreso, 5 m. S., 128 6, 229 2, xi.24.63, JRZ (NMSU).

Laccophilus vacaensis chihuahuae, new subspecies
(Figs. 146, 149, 152, 312)

DESCRIPTION AND DIAGNOSIS. — L. v. chihuahuae differs from v.
vacaensis in its lighter color, less extensive elytral pattern, smaller size, highly

JAMES R. ZIMMERMAN 161

modified female epipleura, and the structure of the male aedeagus. Yellow or
yellowish-brown predominates as the principal base color of chihuahuae instead
of the reddish-brown that is much more apparent in vacaensis. The irroration
of chihuahuae is less intense with the individual dots smaller and more dis-
cretely separated from one another. The average size of chihuahuae is about
0.3 to 0.4 mm shorter than vacaensis. The width of the pronotum exceeds the
length 2.50 times in the latter and 2.57 times in the former (n=8 for each
race). The female epipleura and male aedeagus are most strikingly different
characters, however. The female epipleuron of chihuahuae almost always has a
flange and an unusual conformation immediately anterior to it. The epipleuron
is modified so that a groove can be distinguished even when the flange is com-
paratively small. This does not appear in vacaensis, but does in v. thermophilus.
The aedeagus of vacaensis has a rather sinuate appearance, while that of chi-
huahuae is comparatively straight except just before the apex where it abruptly
narrows to a curving bent tip. Also, if cut in cross-section at about half its
length, the aedeagus would have a decidedly triangular outline. This is only
suggested in vacaensis.

The aedeagus is the only reliable way to separate chihuahuae and ther-
mophilus, although the latter seems to be slightly larger (Table 21).

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L. v. chihuahuae ranges from the hill country of central Texas
through west Texas, New Mexico, and Cochise County, Arizona (fig.
15). I have collected it only in short-lived ephemeral ponds that
occur after the summer rains in the flats between the separated moun-
tain ranges of New Mexico, Arizona, and west Texas. This ephem-
eral condition seems to be the reason this race is so seldom collected.
It does occur in the lower reaches of some mountain streams, however.

MATERIAL EXAMINED

Holotype male, allotype female, and a male paratype with the following
locality data are in the United States National Museum: Boquillas, Big Bend
National Park, Brewster County, Texas, viii.2.61, J. R. Zimmerman. One male
and two female paratypes with the following data are in the California Acad-
emy of Sciences, San Francisco: 1 m. S. of Animas, Hidalgo County, New
Mexico, vii.31.65, H. B. Leech; one male paratype, 5 miles N. of Tombstone,
Cochise County, Arizona, ix.6.61, J. R. Zimmerman, and one female, 10 m. E.
of Marathon, Brewster County, Texas, viii.1.61, J. R. Zimmerman, in the Uni-
versity of Michigan Museum of Zoology, Ann Arbor; and another female para-
type from the same locality is also in the United States National Museum.

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Chiraca-
hua Mtns., Cave Ck. Can., 1 2, ix.13-14.52, B. Malkin (FM). MHuachucha
City, 2 6, 3 2, ix.6.61, JRZ (NMSU). Rodeo (N.M.), 2 m. NW., 2 6, 2 2,

MEM. AMER. ENT. SOC., 26

162 THE GENUS LACCOPHILUS

ix.5.61, JRZ (NMSU). Tombstone, 5 m.S., 1 6, 1 2, ix.6.61, JRZ (NMSU).
NEW MEXICO. Harding County. Mosquero, 10 m. E., 1 2, x.23.65, A. H.
Smith (NMSU). Hidalgo County. Rodeo, 1 2, viii.31.59, H. E. Evans
(CNL); 12 m.N., 1 6,2 2, ix.5.61, JIRZ (NMSU). TEXAS. Bexar County.
Leon Ck., 1 6, 3 2, x.11.52, B. J. Adelson (CAS). Brewster County. Bo-
quillas, 1 2, vii.7.48, C. & P. Vaurie (AMNH); 3 6, 4 2, vili.2.61, JRZ
(NMSU). Marathon, 10 m. E., 4 2, viii.1.61, JRZ (NMSU). Burnett County.
1 $6, 8 2, Hubbard and Schwarz (USNM). Culberson County. Nickel Ck.
Sta., 2.5 m. E., 2 9, ix.2.52, B. Malkin (FM). Lee County. Fedor, 1 2
(CNG). Webb County. Laredo, 2 ¢, v.20-24.48, W. Nutting and F. Werner
(ARI).

Laccophilus vacaensis thermophilus, new subspecies (Fig. 154)

DESCRIPTION AND DIAGNOSIS. — L. v. thermophilus differs from L. v.
vacaensis in much the same way that chihuahuae does; i.e., in color, pattern,
size, and sexual characters. It appears to differ from chihuahuae only in the
structure of the male aedeagus and perhaps in being slightly larger. The tip of
the aedeagus is blunter than that of chihuahuae. The relative length of the
pronotum is almost the same as vacaensis (2.52 times as wide as long). This
race appears to occur all around the Gulf of California; but, as yet, the records
are too scattered for verification. It apparently prefers the same kind of habitat
as chihuahuae, but in a hotter climate.

VARIATION. — The specimens from Baja California average
much larger than the ones from Sonora. However, there are only a
total of six specimens available from the latter area. The standard
deviations are high enough to suggest that even that material is mixed.
Since all three races of vacaensis might occur in that area, it is entirely
possible that there is some intergradation represented in the Sonoran
specimens determined as thermophilus.

MATERIAL EXAMINED

Holotype male, allotype, and fifteen paratypes with the following data from
8.2 m. W. La Pas on Hwy. Sur No. 1, Baja California Sur, xii.13.58; H. B.
Leech are in the California Academy of Sciences, San Francisco.

UNITED STATES OF AMERICA. — ARIZONA. Pima County. Arivaca,
Arivaca Ck., 1 6, vii.31.52, H. B. Leech (CAS).

MEXICO. — SINALOA. Mazatlan, 1 9, xii.11.62, JRZ (NMSU). SONORA.
Pitiquito, 2 m. E., 1 6, viii.1.50, J. Figg-Hoblyn (CAS). Desemboque, 1 2°,
ix.1-10.53; Navojoa, 1 2, ix.26.53, B. Malkin (CAS). Guaymas, 2 2, ix.29.23,
W. M. Mann (USNM).

Laccophilus spangleri, new species (Figs. 155-162, 315)
DIAGNOSIS. — Laccophilus spangleri can best be identified by the follow-

JAMES R. ZIMMERMAN 163

j
\2 LACCOPHILUS V. VACAENSIS e

: ©] LACCOPHILUS Vv. CHIHUAHUAE 0

LACCOPHILUS V. THERMOPHILUS 0

LACCOPHILUS SPANGLERI 9

Figure 15. Distribution of Laccophilus vacaensis and L. spangleri.

ing combination of characters: irrorated pattern, lack of file in males or females,
rakelike ovipositor, and the fifth front tarsal segment less than twice as long as
the corresponding fourth. The rakelike ovipositor and lack of a file separates
spangleri from most irrorated species, but it is easily confused with vacaensis
and peregrinus. Its size is intermediate and overlaps with both species. The
front tarsal segment of the males of vacaensis is more than twice as long as the
corresponding fourth and helps to separate males, but females cannot be sepa-
rated with complete assurance. Irroration in spangleri tends to be darker with
more fusion of the individual dots. In specimens from Mexico the “fingers”
that form the anterior part of the elytral pattern are more clearly defined and
do not tend to fuse anteriorly, while in v. vacaensis there is considerable fusion
with discrete “fingers” not really apparent. South of Mexica, this is not a relia-
ble difference. The darkened area on the head and pronotum is more apparent
in spangleri than in vacaensis.

L. proximus and L. fuscipennis, which are sympatric with spangleri in Vera-
cruz and have similar patterns and color, have metacoxal files in the males and
sawlike ovipositors in the females. L. peregrinus peregrinus has a similar aede-
agus and ovipositor; but in Veracruz the elytral pattern is more uniform than

MEM. AMER. ENT. SOC., 26

164 THE GENUS LACCOPHILUS

spangleri, and the average size is about 0.3 to 0.5 mm shorter (Tables 22, 24).

DESCRIPTION. — Medium (length, 4.1 to 4.8 mm; width, 2.3 to 2.7 mm),
brown, irrorated species; metacoxal file absent; prosternal process short; ovi-
positor rakelike. COLOR. Head: generally pale brownish-yellow tinged with
ted above and beneath except for some dark reddish-brown on the occiput
between the eyes. Pronotum: pale yellowish-brown except anteriorly between
the eyes; here darker color matches the darkened area of the head; some of the
darker color trails weakly across the disc and the apex may be translucently
reddish-brown. Elytra: heavily irrorated, reddish-brown pattern on yellow back-
ground; considerable coalescence of dots especially around margins of pattern;
epipleura pale anteriorly and reddish-brown posteriorly. Tergite VIII: basally
dark brown, but posterior half light yellowish-brown. Venter: variably light
yellowish-brown with reddish tinge; front two pair of legs usually lighter than
Test; sutures, coxal bases usually darkened. Genitalia: about the same color as
venter. ANATOMY. Méicroreticulation: clearly double on much of the head
and pronotum, but only weakly so on the elytra, especially in females; surface
shining. Head: supraclypeal seam arching slightly upward at center above
margin. Pronotum: WH/PW, 0.70; LP/PW, 0.40. Elytra: slight truncation
of apices; female epipleura without flange. Venter: prosternal process with a
median crest that nearly attains the anterior prosternal margin; postcoxal proc-
esses rounded, laterally projecting slightly posterior to the midline; last visible
segment of males slightly truncated with produced apex and with an asym-
metrical median crest; a small tuberosity near the left hind margin of the next
to last ventral segment; last female segment not truncated, but nearly triangular;
marginate groove on either side of the apex; both sexes with numerous setiger-
ous punctures. Legs: pro- and mesotarsi enlarged in a dorsoventral plane; fifth
tarsal segment of both legs about one and three-quarters as long as the corre-
sponding fourth; palettes easily visible at 20 power magnification; profemoral
setae (6 or 7) smaller and shorter than mesofemoral ones (6 to 8). Genitalia:
oval plate with produced acuminate tip; its ventral crest curving slightly to the
left as it extends anteriorly to near front margin; numerous raised lines of
varying thickness on either side of the crest; aedeagus curving without notice-
able angle; unevenly tapered to small knob at apex; ovipositor with about eight
pairs of rakelike teeth.

VARIATION. — The elytral irroration varies in intensity and in
the degree to which the dots tend to coalesce. In some specimens
from Veracruz, the color is almost completely suffused with little evi-
dence of irroration. South of Mexico there is much more spreading
and fusing of the “fingers” so that the most anterior part of the pattern
has a more solid appearance similar to that of vacaensis.

Females are slightly larger than males (Table 13, fig. 16), but
there is almost complete overlap. There is no clear geographic trend
in overall size, as seen by the similarity of means for the samples from
La Tinaja, Veracruz, and from San Benito, Nicaragua.

165

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

166 THE GENUS LACCOPHILUS

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This species has been taken from several localities in northern and
central Veracruz, from three localities on the south side of the Isthmus
of Tehuantepec, and in Honduras and Nicaragua (fig. 15). Addi-
tional collecting will probably show that it cccurs in most of the coastal
regions of southern Mexico and Central America. It is a tropical low-
land species. It seems to prefer temporary pools that are frequently
subject to high surface temperatures, and that are located in clay soils.

MATERIAL EXAMINED

Holotype male, allotype female, five male, and five female paratypes with
the following locality data are in the United States National Museum; Tehu-
antepec, Oaxaca, ix.2.63, J. R. Zimmerman. One male and one female para-
type with the same locality data are also in each of the following collections:
California Academy of Sciences, San Francisco, the University of Michigan
Museum of Zoology, Ann Arbor, in the Departmento de Entomologia, Labora-
torio de Sanidad Vegetal, Coyoacan, D. F., Mexico.

HONDURAS. — San Marcos Colon, 3 ¢, 2 9, vii.28.65, P. J. Spangler (USNM).

MEXICO. — OAXACA. Juchitan, 20 km. N., 3 6, 2 2, ix.7.64; 3 m. E., 8 4,
33 2, ix.7.64; Tehuantepec, 26 3, 23 9, ix.2.63, JRZ (NMSU). VERACRUZ.
Cuitlahuac, 1 ¢, viii.25.62; near La Tinaja, 10 6, 8 9, viii.25.62; Paso de
Ovejas, 10 8, 13 9, viii.27.62, JRZ (NMSU). Poza Rica, 2 6, 1 2, viii.27.65;
Tantoyuca, 15 m. SE., 8 6, 12 2, viii.28.65, P. J. Spangler (USNM).

NICARAGUA. — Esteli, 9 m. N., 14 6, 9 @, vii.10.65; La Trinidad, 1 @,
vil.27.65, P. J. Spangler; Rivas, 1 6, vii.31.67, O. S. Flint; San Benito, 13 m. N.,
36 6, 28 2, vii.11.65; Somoto, 3 6, 5 9, vii.11.65, P. J. Spangler (USNM).

LACCOPHILUS PEREGRINUS

This is a highly variable, polytypic species that ranges from Sina-
loa and Veracruz, south to Panama, and probably into South America.
It is composed of two races that are roughly separated by the Isthmus
of Tehuantepec. The northern race, peregrinus, has two allopatric
populations, which conceivably could be split even more into separate
races; but lacking any character that could be demonstrated as con-
sistently different between the two, they were retained as one. Vera-
cruz populations have enough elytral pattern variation that some indi-
viduals with an irrorated pattern could readily be included in the west
coast population. Others, however, have almost uniformly reddish-
brown elytra and, hence, look quite different from the irrorated ones.
The data from measurements are too incomplete to show much differ-
ence in the two populations and, in fact, show a close similarity when

JAMES R. ZIMMERMAN 167

spangleri

v. vacaensis

NUMBERS

4.0 4.2 4.4 4.6 4.8 5.0 5.2
YOvAL EEN Girt (oath)

Figure 16. Histograms of length in Laccophilus v. vacaensis, L. spangleri,
and L. fascipennis. Males are shown crosshatched; females, stippled.

MEM. AMER. ENT. SOC., 26

168 THE GENUS LACCOPHILUS

compared to the values obtained for variabilis. The structure of the
aedeagus supports retaining them as a single race, also.

The southern race, variabilis, has an even more heterogenous
appearance than peregrinus. It varies from a black and yellow varie-
gated form to one with nearly uniformly black elytra to one with a
pattern which is found in typical peregrinus. The variation occurs
in a north-south direction, so that the unlikely situation exists that
individuals from Panama superficially most resemble those from Sina-
loa. The quantitative data, however, show enough uniformity to sug-
gest no great separation or differentiation of the different variabilis
populations.

In this species elytral pattern has been less reliable for the recog-
nition of intergrades than comparison of male aedeagi. In other
species, L. maculosus and L. fasciatus for example, the pattern has
frequently manifested intergrade features while the aedeagus did not.
Only in a few instances were individual males found that had an
aedeagus that clearly represented the intermediate condition. In L.
peregrinus in Chiapas both races are represented as judged by elytral
patterns. The southern race was taken in Ixtapa and nearby Chiapa
de Corzo and the northern race in Comitan, which is about 90 miles
closer to Guatemala. Other specimens of the southern variabilis were
taken from three other localities in Oaxaca. None showed any inter-
gradation in elytral pattern; but the males have unmistakenly inter-
mediate aedeagi, and the aedeagi of the various Chiapas males are
virtually identical.

Additional evidence for intergradation is found in the quantita-
tive data.

The mean values for all four measurements for the Ixtapa and
Chiapa de Corzo females are larger than for any of the other variabilis
populations and approaches the value for typical peregrinus females
(Table 14). In figure 18, 13 of the 18 largest individuals are females
from Chiapas. Note that these females lie at about the mean value
for p. peregrinus females (and males). This suggests that the larger
size is due to gene flow from the northern race.

Laccophilus peregrinus, new species (Figs. 163-172)

DESCRIPTION. — Small (length, 3.3 to 4.6 mm; width, 1.9 to 2.6 mm)
species; elytral pattern highly variable from brown to reddish-brown, black, or
yellow and black, irrorated, unicolorous, or variegated; metacoxal file absent;
prosternal process short; ovipositor rakelike. COLOR. Head: yellow darken-

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

170 THE GENUS LACCOPHILUS

ing to brown or reddish-brown on the occiput between the eyes. Pronotum:
pale brownish-yellow except for prominent dark reddish-brown or brown on the
anterior margin between the eyes and a somewhat less distinct area at the apex.
Elytra: pattern varying from irregularly irrorated brown on a pale yellowish-
brown background to one that is almost entirely unicolorous reddish-brown or
very dark brown or black; or is variegated with yellow and dark brown or
black; epipleura pale anteriorly and pale or dark posteriorly. Tergite VII:
basally dark brown, but lightening to yellowish-brown along the hind margin.
Venter: pro- and mesolegs pale reddish-yellow brown, remainder darkening to
medium shades of the same color; abdominal sternites usually darker than
thorax and metacoxal plates. Genitalia: reddish- or yellowish-brown. ANAT-
OMY. Microreticulation: double on head, pronotum, and elytra. Head: supra-
clypeal seam arching slightly upward above the margin. Pronotum: WH/PW,
0.71; LP/PW, 0.38 to 0.39. Elytra: somewhat attenuated, truncation barely
perceptible; female epipleura usually with a flange in p. peregrinus, but almost
never in p. variabilis; crease apparent immediately above the female epipleura
and extends about one-third the distance along the elytron. Venter: prosternal
process with well-defined crest extending from apex to level of line drawn across
the front margins of procoxal cavities; postcoxal processes rounded and laterally
projecting slightly beyond the midline; last visible segment not truncated in
either sex; rounded in males with very slight production of the apex in males
and subtriangular in outline in females; males with asymmetrical curving crest;
females with marginate groove extending from anterior lateral margin of seg-
ment to near apex; scattered setigerous punctures accumulating toward hind
margin in both sexes. Legs: pro- and mesotarsi enlarged in a dorsoventral
plane; fifth tarsal segments on both pair of front legs about twice as long as
corresponding fourth segment; palettes easily visible at 20 power magnification;
profemoral setae (5 or 6) much finer and shorter than the mesofemoral ones
(5 or 6). Genitalia: oval plate with acuminate produced tip; median crest of
plate extending anteriorly with little or no curvature; about three or four raised
lines on the left side of crest and about twice that number on the right; aedeagus
with strong bend at about half its length and another lesser one subbasally; right
paramere smaller than left; ovipositor with four pair of long wide-spaced distal
teeth and about five more smaller proximal ones.

Laccophilus peregrinus peregrinus, new subspecies (Figs. 316, 317)

DESCRIPTION AND DIAGNOSIS. — Elytral pattern and aedeagus pro-
vide the only reliable clue for identification of L. p. peregrinus within the group
of species that lack a metacoxal file and have a rakelike ovipositor. The size
ranges from a length of 3.7 to 4.6 mm and width of 2.1 to 2.6 mm. Other
species with a similar irrorated pattern are L. spangleri, L. vacaensis and L.
proximus. It is sympatric with all of them. L. proximus has no pronotal dark-
ening, has a metacoxal file, and has a sawlike ovipositor. The males of vaca-
ensis have the fifth tarsal segment on both pair of front legs well over twice
the length of the corresponding fourth, while it is only twice the length in

JAMES R. ZIMMERMAN 171

peregrinus. The microreticulation of the elytra of vacaensis is single and gives
a more pebbled appearance than would be seen in peregrinus. The pronotal
anterior dark area is much more prominent in peregrinus than in vacaensis also.
The crease above the epipleura is more pronounced and longer in vacaensis
females than in peregrinus females. Both vacaensis and peregrinus have the
epipleura often expanded with a flange, however. L. vacaensis (except chihua-
huae) averages more than a half millimeter larger than peregrinus.

L. spangleri can often be very similar in appearance to peregrinus; but, usu-
ally, there is sufficient difference in the details of the elytral pattern of spangleri
to permit separation from the irrorated pattern of peregrinus. If that of pere-
grinus is the suffused, nearly unicolorous type, then there should be no difficulty.
Comparison of male genitalia may be necessary, however. There appears to be
no flange developed on the female epipleura of spangleri. It is a larger species
and averages up to half a millimeter longer.

L. huastecus bears a strong resemblance to L. p. peregrinus, even though it
has a pattern that should more properly be considered suffused than irrorated;
but with the brown on the front of the head and the pebbled appearance of the
elytra, there should be no difficulty in separation.

L. p. peregrinus differs from L. p. variabilis in the structure of the male
aedeagus, in the color and pattern of the elytra, in its generally larger size, and
in the frequent presence of a flange on the female epipleura. The aedeagus is
blunter and heavier in peregrinus. The elytra are more reddish-brown than brown
or black and are not variegated. The length of peregrinus ranges from 3.7 to
4.6 mm as compared to 3.3 to 4.2 mm in variabilis. The average length of
peregrinus is about 4.1 mm and that of variabilis about 3.7 to 3.8 mm.

VARIATION. — The principal pattern variation is occurrence of
both an irrorated pattern and a suffused, nearly unicolorous one. All
degrees of intermediacy are present. Even when there is no trace of
irroration, it is still possible to distinguish the areas that are normally
pale yellow-brown in the irrorated pattern. This variation is probably
geographical since it seems to be most common in southern Veracruz
and Chiapas; but without more information, it is not possible to decide
whether there is a clearly demonstrable intergrade zone or whether the
situation is clinal.

Females tend to be larger than males (Table 14, fig. 18), but the
overlap is great enough that, in Nayarit, the mean values were larger
for males than for females and, in Veracruz, the means were virtually
the same. There is no clear difference in the size of populations from
the west coast versus those from Veracruz. The small sample from
Chiapas gives about the same values.

MEM. AMER. ENT. SOC., 26

172 THE GENUS LACCOPHILUS

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— The race has been taken at several localities on or near the coastal
plain in Sinaloa and Nayarit. The northernmost record is from Ma-
zatlan, and the southernmost one is San Blas, Nayarit; but it undoubt-
edly occurs to the north and south of that area. It occurs throughout
Veracruz at lower elevations (fig. 17). I have most frequently taken
it in temporary ponds with clay bottom and grassy margins.

MATERIAL EXAMINED

Holotype male, allotype female, and two male and female paratypes from
4 miles west of Esquinapa, Sinaloa, iv.12.68, J. R. Zimmerman, are in the
United States National Museum. Other paratypes include a male and female
from Poza Rica, Ruinas El Tajin, Veracruz, viii.27.65, P. J. Spangler, United
States National Museum; a male from Mazatlan, Sinaloa, viii.14.60, Arnaud,
Ross, and Rentz, California Academy of Sciences, San Francisco, California;
and a male and female with the same locality data as the holotype are in the
University of Michigan Museum of Zoology, Ann Arbor, and in the Depart-
mento de Entomologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F.,
Mexico.

MEXICO. — CHIAPAS. Comitan, 2 é, 4 2, viii.30.63, JRZ (NMSU). NAYA-
RIT. Acaponeta, 12 m. S., 1 2, vii.31.62; San Blas, 5 m. E., 10 6, 16 92,
vii.31.62, JRZ (NMSU). SINALOA. Mazatlan, 2 6, vili.14.60, Arnaud, Ross,
Rentz (CAS); 1 9, viii.1.62; 4 6, 2 9, xii.11.62; Esquinapa, 4 m. W., 10 é,
8 2, iv.12.68, JRZ (NMSU). VERACRUZ. Cuitlahuac, 3 2, vili.25.62; Ja-
lapa, 10 m. E., 6 6, 11 Q, viii.27.62; near Garro, 3 2, ix.8.64; near La Tinaja,
12 2, 8 @, vili.25.62; Paso del Toro, 6 6, 17 9, viii.25.62; 15 km. W., 1 2,
ix.5.64, JRZ (NMSU). Poza Rica, 1 6,1 2, viii.27.65, P. J. Spangler (USNM).
Santiago Tuxtla, 10 km. S., 1 6, 1 @, 1x.8.64, JRZ (NMSU).

Laccophilus peregrinus variabilis, new subspecies (Figs. 318, 319)

DIAGNOSIS AND DESCRIPTION. — Elytral pattern is the best way to
distinguish L. p. variabilis. The yellow and black, or nearly black, variegated
pattern, or the almost uniform elytra of the same dark color are unique among
North American Laccophilus. The irrorated pattern of Panamanian variabilis
is quite similar to that of p. peregrinus in general appearance but differs in de-
tails of outline and in being more extensive. This latter pattern might be mis-
taken for spangleri, vacaensis, huastecus, and proximus, but variabilis can be
separated in the same fashion as discussed for peregrinus. Some specimens of
p. peregrinus have almost uniformly pigmented elytra, but they are reddish-
brown. The aedeagus of variabilis is finer and is more tapered than in the
nominate race. It is a smaller race; and the female almost never, if ever, has
a flange on the female epipleuron as in p. peregrinus.

JAMES R. ZIMMERMAN 173

VARIATION. — The problem in variabilis is similar to that in
typical peregrinus in that there is considerable variation in elytral
pattern and the variation is geographical. Specimens from the more
northern parts of the range are variegated, those from the central part
tend to be unicolorous, and some from Panama are irrorated. Not
enough is known, as yet, to delineate any separate races within the
assemblage. There is also some change in the aedeagus from north
to south.

Except for being smaller, the same statements for peregrinus apply
to variabilis. There appears to be no geographical trend indicated by
the data from measurements.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L. p. variabilis ranges from Oaxaca to Panama (fig. 17). While
more frequently taken in coastal lowlands, it is also found at over
5000 feet in Ixtapa, Chiapas. It is in temporary ponds with clay bot-
toms, which have a considerable amount of emergent grass and sedges
around the margins.

MATERIAL EXAMINED

Holotype male, and allotype female, from 13 m. N. San Benito, Nicaragua,
vii.11.65, P. J. Spangler; paratypes as follows: six males and six females, same
data as holotype, USNM; one male and one female, same data, in each of the
following collections: California Academy of Sciences, San Francisco; Univer-
sity of Michigan Museum of Zoology, Ann Arbor; and Departmento de Ento-
mologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F., Mexico.

MEXICO. — OAXACA. Juchitan, 3 m.E., 1 2, ix.7.64, JRZ (NMSU). Oaxaca,
146, 1 2, vi.29.57, D. R. Lauck (USNM). Tapanatepec, 3 6, 2 2, ix.1.63,
JRZ (NMSU). CHIAPAS. Chiapa de Corzo, 6 4,9 9, ix.1.63; Ixtapa, 7 ¢,
5 2, viii.31.63, JRZ (NMSU).

COSTA RICA. —La Cruz, 16 m. S., 50 6, 50 9, vii.25.63; 1 2, vii.13.65;
Liberia, 15 m. SW., 7 6, 8 9, vii.24.65; Taboga Agr. Exp. Sta., 55 ¢, 24 9,
vi.27.67, P. J. Spangler (USNM).

EL SALVADOR. — La Union, 15 m. SW., 1 6, 1 2, vii.31.65, P. J. Spangler
(USNM).

HONDURAS. — San Marcus Colon, 1 2, vii.28.65, P. J. Spangler (USNM).

NICARAGUA. — La Trinidad, 6 6, 2 2, vii.27.65; Rivas, 10 m. N., 1 4,
vii.11.65; San Benito, 13 m. N., 63 6, 51 Q, vii.11.65, P. J. Spangler (USNM).

PANAMA. — Anton, 5.3 m. E., 19 ¢, 7 2, vi——.67; Algarrobos, 9 m. W.,
26,2 2, vii.5.67, P. J. Spangler (USNM).

MEM. AMER. ENT. SOC., 26

174 THE GENUS LACCOPHILUS

LACCOPHILUS P. PEREGRINUS
LACCOPHILUS P VARIABILIS
LACCOPHILUS HUASTECUS

LACCOPHILUS OVATUS ZAPOTECUS

Figure 17. Distribution of Laccophilus peregrinus, L. huastecus, and L.
ovatus zapotecus.

Laccophilus huastecus, new species (Figs. 183-191, 320)

DIAGNOSIS, — L. huastecus belongs to the group of species that have no
metacoxal file, have a rakelike ovipositor, and have the elytral microreticulation
single. It especially resembles L. p. peregrinus and L. spangleri with which it is
sympatric in Veracruz. This species can be separated from all other North
American species of Laccophilus, however, by the brown color present on the
front of the head. It can also be distinguished from peregrinus by elytral pat-
tern and male aedeagus. L. p. peregrinus has an irrorated irregular pattern, or
suffused, nearly unicolorous reddish-brown elytra, while Ahuastecus has a suf-
fused irregular brown pattern with three distinct sub-basal elongated clear areas.
It is possible that huastecus, as other species in the group mentioned above, may
have the pattern expressed in irrorations as well as being suffused. In the three
specimens upon which this description is based, one had a suggestion of irrora-
tions on the lighter portions of the pattern. The aedeagus of huastecus is one
that is gradually curved to a tapered tip, while that of peregrinus has a strong
bend at about half its length and a blunt tip. The single female of huastecus
I have seen had no suggestion of an epipleural flange, while almost all pere-
grinus females do have a flange.

JAMES R. ZIMMERMAN 175

peregrinus

variabilis

NSUSMe 8 ERs

4.4

TiO AGE EVE NOG HS (mim)

Figure 18. Histograms of length of Laccophilus peregrinus. Males are
shown crosshatched; females, stippled.

MEM. AMER. ENT. SOC., 26

176 THE GENUS LACCOPHILUS

L. huastecus is smaller than spangleri. The former is about 4.0 mm long
and spangleri usually averages more than 4.4 mm. The elytral patterns of the
two are similar, but that of spangleri is less extensive, especially subbasally, and
has a much greater tendency toward irroration.

L. huastecus also resembles peregrinus variabilis anatomically; but the ely-
tral patterns and male aedeagi can be used to separate them, in addition to the
lack of brown on the front of the head of p. variabilis.

DESCRIPTION. — Small (length, 3.9 to 4.2 mm; width, 2.0 to 2.2 mm),
dark brown and yellow species; metacoxal file absent; prosternal process short;
ovipositor rakelike. COLOR. Head: yellowish-brown on front, dark brown on
back between the eyes, darker immediately medial to eyes, but paler in center
of head. Pronotum: pale yellow and very lightly tinged with reddish-brown
except for dark brown on front margin between the eyes and at apex. Elytra:
dark brown, irregular, suffused pattern on a yellow background; some sugges-
tion of irrorations in lighter areas; pattern with three prominent elongated clear
areas near base; epipleura pale anteriorly, darkened posteriorly. Tergite VIII:
pale yellowish-brown distally but dark brown at base. Venter: pro- and meso-
legs pale yellow; metacoxal plates darkened slightly with brown; last three
abdominal sternites darkened to brown, while basal two remain pale brownish-
yellow; hind legs about the color of basal sternites. Genitalia: yellowish-brown
tinged with red. ANATOMY. Méicroreticulation: faintly double on head and
pronotum, but single on the elytra. Head: supraclypeal seam arching upward
at middle above margin. Pronotum: WH/PW, 0.69; LP/PW, 0.39. Elytra:
truncation barely perceptible; somewhat attenuated. Venter: prosternal process
with well-defined crest in apical third; postcoxal processes rounded laterally and
projecting slightly posterior to midline; last visible segment not truncated, but
rounded with slightly produced apex in males and subtriangular in females;
males with asymmetrical curving median crest; females with marginate groove
extending from anterior lateral margin of segment to near apex; both sexes with
scattered setigerous punctures accumulating toward posterior margin. Legs:
pro- and mesotarsi enlarged in a dorsoventral plane; fifth tarsal segment of pro-
legs slightly more than twice as long as corresponding fourth; profemoral setae
(6 to 7) much finer and shorter than mesofemoral ones (6?). Genitalia: oval
plate with triangular acuminate produced tip and median crest that extends an-
teriorly with slight leftward curvature; about six or eight raised lines on either
side of the crest; aedeagus curving gradually to tapered tip; slight twist just
before apex; right paramere smaller than left, apex blunted; ovipositor with five
pair of wide-spaced distal teeth and about five pair of much smaller sawlike
ones proximally.

VARIATION. — The single female examined is larger than both
males. The elytral patterns of the three are much alike and differ
primarily in color intensity.

GEOGRAPHICAL RANGE. — This species is known from two
localities in the northern third of Veracruz. One male was taken from

JAMES R. ZIMMERMAN i7/7/

the ruins at E] Tajin near Poza Rica, and a male and a female were
taken fifteen miles southeast of Tantoyuca (fig. 17).

MATERIAL EXAMINED

Holotype male, 15 m. SE. Tantoyuca, Veracruz (label gives Pue., but it has
to be in Veracruz), Mexico, viii.28.65, P. J. Spangler; length, 4.10 mm; elytral
length, 3.10 mm; width, 2.16 mm; pronotal width, 1.75 mm; PW/EL, 0.564.
Allotype female, same locality data as male; 4.21 mm; 3.29 mm; 2.26 mm; 1.84
mm; 0.559; paratype, male, Poza Rica, Veracruz, viii.27.65, P. J. Spangler;
3.88 mm; 3.02 mm; 2.05 mm; 1.65 mm; 0.546.

Laccophilus raitti, new species (Figs. 173-182, 311)

DIAGNOSIS. — L. raitti belongs to the group of species which lack a meta-
coxal file, have a rakelike ovipositor, and vary in whether they are irrorated or
not. L. raitti would generally not be considered irrorated; but, in one female,
the pigmentation is sufficiently weakened to make it appear that there is some
irroration with suffusion of color between dots. The lack of a file, and the
elongated front tarsi in males, the widespread teeth on the ovipositor and the
long crease above the epipleura in females, and the elytral pattern in both sexes
should separate raitti from other Laccophilus, except vacaensis. L. raitti is very
similar anatomically to vacaensis, but the latter has the male front tarsi com-
paratively longer, and has a slightly different conformation of the aedeagus;
also, the crease above the epipleura on the elytra does not extend as far pos-
teriorly. If the typical elytral pattern of raitti is present, however, there will be
no difficulty in separating the two species. Some L. peregrinus have elytral pat-
terns similar to that of raitti, but the race with which it is sympatric is irrorated
and that difference is enough to permit field separation. The front tarsi of
males are shorter, and the crease is weaker in females of peregrinus; and it is a
smaller species. L. spangleri also belongs to this group, but it differs from raitti
in the ways just enumerated for peregrinus.

DESCRIPTION. — Medium (length, 4.8 mm; width, 5.2 mm) brown and
yellow variegated, marmorated, or irrorated species without metacoxal file; pro-
sternal process short; ovipositor rakelike. COLOR. Head: generally pale
brownish-yellow above and beneath, but with reddish-brown occiput between
the eyes. Pronotum: pale brownish-yellow except for reddish-brown anterior
margin between the eyes and confluent with dark color on head; also darkened
less intensely at apex. Elytra: usually variegated with dark reddish-brown on
a pale brownish-yellow background, dark area usually marmorated, but may be
faintly irrorated with considerable suffusion of color between the dots (one
specimen in a total of 14); epipleura pale anteriorly and translucent reddish-
brown in narrowed posterior part. Tergite VIII: pale yellowish-brown. Venter:
variably pale yellowish-brown or light reddish-brown; pro- and mesosternum
and their legs usually lightest; abdominal segments and hind legs darkest. Geni-
talia: about the same as venter, except that the right paramere tends to be dark

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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JAMES R. ZIMMERMAN 179

reddish-brown. ANATOMY. Microreticulation: head and pronotum double
over much of their surfaces, but elytra weakly double in males and scarcely or
not at all in females which results in a pebbling similar to females of vacaensis;
inner mesh cellules of head well-defined within the larger matrix. Head: supra-
clypeal seam arching slightly upward at middle above margin. Pronotum:
WH/PW, 0.69; LP/PW, 0.39. Elytra: apices slightly truncated in both sexes;
in females well-defined supra-epipleural ridge extending along most of the ely-
tral length (apparent, but much weaker in males); female epipleura without
flange (none in seven specimens). Venter: prosternal process with well-defined
median crest in distal three-quarters; postcoxal processes rounded and laterally
projecting posterior to the midline; last visible segment of both sexes not trun-
cated, but rounded with a produced apex in males and subtriangular in females;
males with an asymmetrical median crest; females with broad, triangular crest
and marginate groove along hind margin; both sexes with a few small rugae and
scattered setigerous punctures. Legs: male pro- and mesotarsi enlarged in a
dorsoventral plane and also slightly dilated laterally; fifth tarsal segment of both
pair of anterior legs twice as long as corresponding fourth; palettes easily visible
at 20 power magnification; profemoral setae (female, 7; male, 8 to 10). Geni-
talia: oval plate with produced pointed apex; its ventral median crest bent to
the left; numerous raised lines on either side of the crest; aedeagus sharply
angled at about one-half its total length and with small knob at tip; right para-
mere with produced apex; ovipositor with five pair of rakelike teeth.

VARIATION. — Among the handful of specimens of this species
that I have seen, the principal pattern variation is that, instead of pig-
ment suffusing evenly over the elytra, there is a strong suggestion of
irroration. Only one specimen out of 14 was clearly irrorated, but
two more were slightly so. The intensity of the pigment varies from
very dark to medium reddish-brown. The non-pigmented area of the
elytra vary also.

Sexes appear to be about the same size with the means of all four
measurements being statistically the same (Table 15). The condition
of the specimens were somewhat poor, however; and better samples
(and larger ones) are needed to make a reliable conclusion.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— L. raitti is known from a single locality and collection. The speci-
mens were taken in a roadside ditch about five miles inland on the
coastal plain near San Blas, Nayarit. The pond was a shallow (no
more than a foot deep at any point) temporary one with mud bottom
and some grass around the margin. The surrounding vegetation was
tropical evergreen forest. Seven other species of Laccophilus were
taken from the same locality.

MEM. AMER. ENT. SOC., 26

180 THE GENUS LACCOPHILUS

MATERIAL EXAMINED

Holotype male, allotype female, and a paratype of each sex with the follow-
ing locality data are in the United States National Museum: 5 m. E. of San Blas,
Nayarit, vii.31.62, J. R. Zimmerman. A paratype of each sex with the same
data are also deposited in the California Academy of Sciences, San Francisco,
the University of Michigan Museum, Ann Arbor, and the Departmento de
Entomologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F., Mexico.

MEXICO. — NAYARIT. San Blas, 5 m. E., 2 ¢, 2 2, vii.31.62, JRZ (NMSU).

LACCOPHILUS QUADRILINEATUS

This is the largest species of North American Laccophilus. It was
described by Horn in 1871 from Texas. It is a highly variable species
and, as interpreted in this paper, has a range that covers most of the
central and south Plains, the Southwest, and much of Mexico. It is a
polytypic species with three races that are, apparently, completely
allopatric from one another. For this reason, it could easily be inter-
preted as three species instead of one. The species is closely related
to L. inagua Young (1963) from the Bahamas. This suggests that
additional collecting should turn up other populations through the
West Indies. It is also very closely related to L. sonorensis n. sp.
from the southwestern United States, Chihuahua, Sonora, and Baja
California.

This species group is sufficiently different in size, shape, pattern
and several anatomical characters from all other North American Lac-
cophilus to pose a problem as to its origin and as to which group of
species it belongs.

DESCRIPTION. — Large (length, 4.9 to 6.7 mm; width, 2.6 to 3.8 mm),
brown and yellow, or black and yellow, variegated, vittate, or marmorated spe-
cies; metacoxal file absent; prosternal process short; ovipositor rakelike. COLOR.
Head: pale brownish-yellow or brownish-red; specimens from southern Mexico
with dark reddish-brown occiput. Pronotum: same color as head, but mayae
with irregular, laterally elongated darker reddish-brown discal mark in some
specimens. Elytra: varying in the different races from a dark blotch with four
anterior extensions on a pale yellow background to generally dark brown with
a few discrete yellow markings; dark color may be solid or marmorated; epi-
pleura always pale anteriorly, but may be dark or pale posteriorly. Tergite
VIII: brown or dark brown at base, but may be pale yellow brown to brown
along the posterior margin. Venter: variable from pale brownish-yellow, espe-
cially the pro- and mesolegs, to frequently dark reddish-brown on the metacoxal
plates and the abdominal sternites. Genitalia: variable from pale brownish-

181

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

182 THE GENUS LACCOPHILUS

yellow to dark reddish-brown. ANATOMY. Microreticulation: weakly double
on head and pronotum with inner cellules nearly as distinct as secondary net-
work; elytra single, and more pebbled, especially females. Head: supraclypeal
seam deeply impressed and closely parallel to the margin. Pronotum: mid-
discal depression longitudinally elongate instead of round as in most of the
North American species of the genus; posterior margin sharply deflected form-
ing a more acute apical angle than is common in the genus; WH/PW, 0.68 to
0.69; LP/PW, 0.41 to 0.42. Elytra: apices somewhat truncated; epipleura of
females with anterior wide portion extending posteriorly as far as the third
abdominal segment instead of the first as in most of the genus; flange present in
tehuanensis females. Venter: prosternal process broad and with raised median
crest; postcoxal processes laterally projecting slightly beyond the midline; last
visible segment in both sexes not truncated; apex produced in male; crest in
both sexes poorly defined; viewed ventrally in both sexes the lateral sides of the
segment slope sharply upward causing the posterior margin to be less rounded
and more angular; apex of females reflected ventrally; weak marginate groove
in females; numerous rugae and setigerous punctures scattered over surface.
Legs: pro- and mesotarsi enlarged in a dorsoventral plane and also slightly
dilated laterally; fifth protarsal segment twice as long as corresponding fourth
and fifth mesotarsal segment more than twice as long as corresponding fourth;
palettes easily visible at 20 power magnification; pro- and mesofemoral margi-
nal setae very long (at least as long as femur is wide); those on profemur
(usually 6) shorter and finer than those on mesofemur (usually 7). Genitalia:
oval plate with elongated tip; its ventral crest distinct only near apex; numerous
fine raised lines on either side of crest; aedeagus wider in distal than basal half,
but narrows near apex to a small slightly bent tip; parameres of nearly equal
length; right one with elongated apex; ovipositor more rakelike than sawlike,
with about 11 pair of teeth.

Laccophilus quadrilineatus quadrilineatus Horn, new status
(Figs. 192-198, 306, 307)

Laccophilus quadrilineatus Horn, 1871, p. 330. Lectotype: female, Academy
of Natural Sciences, Philadelphia, no. 2962, Texas; Crotch, 1873, p. 400;
Sharp, 1882a, p. 294; Leng, 1920, p. 77; Zimmermann, 1920, p. 25; Leech
and Chandler, 1956; Young, 1963, p. 1.

DIAGNOSIS. — Size, shape, and elytral pattern make this one of the most
distinctive members of Laccophilus. The same statement applies to other races
of quadrilineatus. In the United States and northern Mexico where this race
occurs, this and L. sonorensis are the only large, non-irrorated species without
a coxal file in the male and without a sawlike ovipositor in the female. Figures
306 and 307 will permit easy separation of the race from other races and simi-
lar species.

NOMENCLATURAL NOTES. —No individual specimen was
selected by Horn from those he examined for his description of the

JAMES R. ZIMMERMAN 183

species. A female lectotype was designated. In this species the mark-
ings and other anatomical features, especially the female epipleura,
make it as reliable for a type as the male. Texas was given as the
locality, which does not seem to include more than one race and no
intergrade zones. The type locality is here restricted to Bexar County,
Texas.

VARIATION. — The degree to which the pattern covers the elytra
varies greatly. The blotch of dark pigment may be extensive, cover-
ing almost half the surface of the elytra or may be reduced to a re-
stricted area without connection with the anterior extensions. The
latter condition is restricted to specimens from southern New Mexico
and Arizona. It may represent a racial difference, but specimens from
California are extremely similar to those from south and central Texas.
At this time it seems better not to describe the variation as new until
the limits and nature of it is better understood.

Females are larger than males (fig. 20). The average length for
females is 5.99 mm and for males 5.72 mm. The largest female is
about 6.4 mm long as compared to the largest male, which is 6.25
mm. The shortest male is about 4.9 mm and the shortest female is
5.4 mm (Table 16).

There is no general geographical trend in size, but the populations
from south and central Texas appear to be somewhat smaller than
those from the rest of the range. The two samples from southwest-
ern New Mexico and Pima County, Arizona, which overlap the range
of L. sonorensis, give mean values for all four measurements that are
as large as any sample of q. quadrilineatus. Hence there is no evi-
dence of intergradation between these two, based upon quantitative
characters.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L. q. quadrilineatus ranges throughout the Plains from northeast-
ern Kansas and east central Colorado south to northern Mexico in
Nuevo Leon, Coahuila, and Chihuahua, and west to Los Angeles,
California (fig. 19). It appears to be limited by woodland on the
east, and by mountains in the south and southwest. It is sympatric
with L. sonorensis from southcentral New Mexico to Blythe, Cali-
fornia. The large gaps in north Texas and southwestern Arizona are
probably due to lack of collecting, since all Laccophilus species from
those areas are poorly represented in collections.

MEM. AMER. ENT. SOC., 26

184 THE GENUS LACCOPHILUS

The species prefers temporary aquatic situations that frequently
have alkaline or salty water with mud bottoms and little or no vege-
tation. In some seasons probably, it is the most common Laccophilus
in stock ponds in the Southwest. While usually not found in the
mountains, it has been taken as high as 8000 feet in Lincoln County,
New Mexico. One of the best localities to collect this species, and
L. sonorensis, is in the heavily alkaline playa near Interstate 10 in
Hidalgo County, New Mexico. Obviously, the species can tolerate
high temperatures and can fly long distances from one drying puddle
to another.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Chiraca-
hua Mtns., Cave Ck. Can., 1 9, ix.13-14.52, B. Malkin (FM). Tombstone,
5 m.N., 44, 1 2, ix.6.61; Huachucha City, 2 2, ix.6.61; Rodeo (N.M.), 2 m.
NW., 5 6, 2 9, ix.5.61, JRZ (NMSU). CALIFORNIA. Imperial County.
Brawley, 1, vili.8.14, J. C. Bradley (CNL). Calexico, 3, vili.24.37; El Centro,
1, —.—.37, H. B. Leech (CAS). County, 1 ¢, viii.27.—, Blaisdell (CAS).
Los Angeles County. Los Angeles, 1 2, vi.16.25 (BERK). Riverside County.
Blythe, 8, vii.10.47, J. W. MacSwain (CAS); 1, viii.6.47, J. W. MacSwain
(BERK). San Bernardino County. Colton, 1, Chittenden (USNM). COLO-
RADO. Kiowa County. Eads, 8 m. S., 2 2, ix.11.63, K. L. McWilliams
(NMSU). Lincoln County. Hugo, 8 m. SE., 2 6, 1 9, ix.11.63, K. L. McWil-
liams (NMSU). KANSAS. Harvey County. County, 1, Popenoe (USNM).
Newton, 5 m. E., 1 2, vii.6.60; near Sedgwick, 1 2, vii.13.60, JRZ (NMSU).
McPherson County. McPherson, 8, viii.29.— (USNM); 3 6, 5 2, vi.20.—,
W. Knaus; 3 6, 5 9, viii.29.—, W. Knaus (AMNH); 1 ¢, vi.4.— (MCZ).
Marion County. Marion, 2.5 m. NW., 1 @, viii.9.60, JRZ (NMSU). Potta-
watomie County. Onaga, 1 (USNM). Reno County. County, 2, ix.21.11,
Blaisdell (CAS). Haven, 8 m. E., 4 6, 1 92, vii.13.60, JRZ (NMSU). Sedg-
wick County. Wichita, 1 ?, JRZ (NMSU). Stafford County. Salt Flats
Area, 2, vii.l1.57, H. & M. Evans (CNL); 1 2, same data (CAS). NEW
MEXICO. Chaves County. Dexter, 5 m. S., 1 9, vil.27.65, A. H. Smith
(NMSU). Dona Ana County. Las Cruces, 1 9, vii.5.61; 1 6, vii.6.61; 4 6,
3 2, vii.10.61, JRZ (NMSU). Jornada Range, Taylor Wells, 19 6, 29 9,
vli.28.63, K. L. McWilliams; 14 ¢, 12 9, vii.20.63, K. L. McWilliams (NMSU).
Mesilla, 1 m. SW., 1 6, vili.1.61, JRZ (NMSU). Guadalupe County. Santa
Rosa, 5 m. NE., 1 6, 1 92, ix.10.63, K. L. McWilliams (NMSU). Hidalgo
County. Animas, 1 m. S., 3 6, 4 9, vii.31.65, H. B. Leech (CAS). Lincoln
County. Alto, 1 @, vii.27.61, JRZ (NMSU). Carrizozo, 13 m. NE., 1 ¢,
3 2, K. L. McWilliams (NMSU). Luna County. Columbus, 1 m. S., 1 @,
vii.17.61, JRZ (NMSU). Deming, 1, —.22.07, Hubbard & Schwarz (USNM).
Hurley, 1 2, vii.17.61, JRZ (NMSU). Otero County. Pinon, 40 m. S., 3 ¢,

JAMES R. ZIMMERMAN 185

3 2, vii.29.63, R. D. Ohmart (NMSU). Socorro County. Near Bingham,
1 6, vii.21.62, R. D. Ohmart (NMSU). Quay County. San Jon, 1 6, 1 2,
ix.10.63, K. L. McWilliams (NMSU). OKLAHOMA. Kay County. Black-
well, 2m. NW., 3 6, vi.30.60; Braman, 2 m. NW., 2 6, 4 9, vi.28.60; 8 m. N.,
1 3,1 9, vi.20.60; Tonkawa, 4 m. NW., 8 4, vi.30.60, JRZ (NMSU). Logan
County. Guthrie, 1, vi.23.11, Kaeber (USNM). Marshall County. 1 (USNM).
Texas County. Hooker, 7 m. SW., 1 6, ix.11.63; Guymon, 1 9, ix.11.63,
K. L. McWilliams (NMSU). TEXAS. Bexar County. Leon Ck., 3 2, 1 2,
x.11.52; 1 ¢, x.12.52; Fort Sam Houston, 2 2, vi.—.52, B. J. Adelson (CAS).
Blanco County. Shovel Mtn., 2 6, 1 9, F. G. Schaupp (AMNH). Brewster
County. Marathon, 22 m. S., 1 6, ix.3.60, L. A. Stange (DAV). Boquillas,
BBNP, 13 6, 17 9, viii.2.61, JRZ (NMSU). Burnet County. County, 3,
Hubbard & Schwarz (USNM). Cameron County. Brownsville, Esperanza
Ranch, 2, viii.14.— (USNM); 1, viii.14.28, L. A. Sayon (USNM); 1, H. S.
Barber (USNM); 3, ix.27.42, E. S. Ross (CAS); 1 (CNL). Port Isabel, 14,
x.20.49, O. Bryant (CAS). Dimmit County. Caterina, 1, vil.7.48, F. Werner
& W. Nutting (ARI). El Paso County. Clint, 1 m. S., 1 2, vii.19.61, JRZ
(NMSU). Hidalgo County. Texas Exp. Sta., light trap, 2 ¢, 1 2, v.29.30;
1, v.30.30; 1, v.20.30, J. C. Gaines (TAM). J. Davis County. Davis Mtns.,
Limpia Canyon, | ¢, 1 9, vii.2.47, B. Malkin (FM). Ft. Davis, 1 6, vii.8.64,
Arnett & Van Tassel (CUA). Kleburg County. Kingsville, 3 (USNM); 7,
C. T. Reed (CNL). Riviera, 3.5 m. N., 1 2, vi.29.61, R. L. Westcott (LACM).
Lee County. 2 (USNM); Fedor, 4 6,4 2 (CNG). Nueces County. Corpus
Christi, 3, vi.10.47, A. Fenyes (CAS). Presidio County. Marfa, 14 m. N.,
3 2, vili.1.61, JRZ (NMSU). Starr County. Rio Grande City, 15 m. NE.,
146, 1 9, 11.3.54, H. F. Howden (USNM). Uvalde County. Uvalde, 1,
v.21.38,S.E.Jones (TAM). Victoria County. Victoria, 1 2, ix.—.— (USNM).
Webb County. Laredo, 1 6, vil.2.47, B. Malkin (FM).

MEXICO. — CHIHUAHUA. Camargo, 1 6, 1 2, vili.8.53, R. & J. Selander
(CAS); 1 m. N., 1 6, vii.25.62, IRZ (NMSU). COAHUILA. Villa Acuna,
San Lorenzo Rnch., 1 ¢, vi.16.38, R. H. Baker (TAM). Ramos Arizpe, 1 ¢,
vii.7.63, JRZ (NMSU). NUEVO LEON. Monterrey, 1 2, x.—.59 (NMSU).
Vallecillo, 3 ¢, 3 2, vi.2-5.51, P. Hurd (BERK).

Laccophilus quadrilineatus tehuanensis, new subspecies (Fig. 309)

DIAGNOSIS AND DESCRIPTION. — L. q. tehuanensis differs from other
subspecies of quadrilineatus in its larger size (Table 17; fig. 19), its nearly black
elytral pattern, the frequent occurrence of a female epipleural flange, and the
shape of the aedeagus. It differs from typical quadrilineatus in that the elytral
pattern is usually more extensive, with the anterior extensions or “fingers”
widening so that they frequently merge instead of remaining discrete. The rest
of the exoskeleton is darker also, especially on the head and venter. The lateral
margins of the metacoxal plates and the area around the bases of the coxae are
dark brown or nearly black. The elytral pattern of mayae is usually more

MEM. AMER. ENT. SOC., 26

186 THE GENUS LACCOPHILUS

extensive, reddish-brown, and somewhat marmorated. The pronotum is rela-
tively longer in tehuanensis than in quadrilineatus or in mayae (2.43 times as
wide as long as compared to 2.33-2.40). The most posterior mesotibial spur
is both relatively and absolutely longer in tehuanensis. It is more than twice
as long as the one next to it, while it is less than twice as long in the other
two subspecies.

VARIATION. — Females average about 0.2 mm longer than
males. The largest female is nearly 6.6 mm long (n= 56) and the
largest male about 6.4 mm (n= 48). The smallest female measured
about 0.1 mm smaller than the smallest male, however (5.6 and 5.7
mm).

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This race is known from three localities on the southern side of the
Isthmus of Tehuantepec and from one specimen taken at Mazatlan,
Sinaloa. It was taken on the coastal lowland in roadside ditch ponds
that had muddy bottoms and highly turbid water. There was no ap-
preciable vegetation except some algal filaments.

MATERIAL EXAMINED

Holotype male, allotype female, four paratypes with the following locality
data are in the United States National Museum. Tehuantepec, Oaxaca, ix.2.63,
J. R. Zimmerman. Two paratypes of each sex with the same data are also in
the California Academy of Sciences, San Francisco, and the Departmento de
Entomologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F., Mexico. A fe-
male paratype from 2 m. W. of Tehuantepec, Oaxaca and a male and female
paratype with same data as the holotype, vi.20.58, J. C. Schaffner, are in the
University of Michigan Museum of Zoology, Ann Arbor, and another female
paratype from Mazatlan, Sinaloa, viii.2.53, C. and P. Vaurie, is in the American
Museum of Natural History, New York.

MEXICO. — OAXACA. Tehuantepec, 46 6, 49 9, ix.2.63; Juchitan, 3 m. E.,
8 6, 10 @, ix.7.63, JIRZ (NMSU).

Laccophilus quadrilineatus mayae, new subspecies (Fig. 310)

DIAGNOSIS AND DESCRIPTION. — L. q. mayae differs from other sub-
species of quadrilineatus in its extensive dark reddish-brown elytral pattern and
in the laterally elongated darkened area on the pronotum. The pattern is lighter
than that of tehuanensis, but the remainder of the exoskeleton has about the
same dark yellowish-brown to brown tones. In all features it is much darker
than quadrilineatus. The proportions of the pronotum are the same as those of
that race (2.33 to 2.40 times wider than long), but relatively shorter than that
of tehuanensis (2.43). The hind margins of the postcoxal processes form
almost a right angle to the midline in mayae — instead of an obtuse angle or

JAMES R. ZIMMERMAN 187

LACCOPHILUS Q.
QUADRILINEATUS e

LACCOPHILUS

SONORENSIS 4

LACCOPHILUS Q.
TEHUANENSIS ©

Figure 19. Distribution of Laccophilus quadrilineatus and L. sonorensis.

an undulating margin as in the other two subspecies. L. q. mayae differs from
L. inagua only in its smaller size (females average 5.80 as compared to 6.75;
n=7), in the less extensive elytral pattern, and in the detailed structure of the
aedeagus. In all other features, the two are extremely similar.

VARIATION. — Females are larger than males (Table 17). The
largest female is about 6.3 mm long and the largest male about 5.8
mm. The smallest female is slightly more than 5.4 mm and the small-
est male is under 5.1 mm (fig. 19). Males may be slightly wider than
females, however. The PW/EL ratios are 0.539 for males and 0.531
for females with no overlapping at the 95 percent confidence interval,
indicating that males may be slightly broader than females.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This subspecies is known from but two localities on the Yucatan
Peninsula. More than a hundred specimens were taken in a single
collection near Progreso, Yucatan, in a roadside puddle that may have

MEM. AMER. ENT. SOC., 26

188 THE GENUS LACCOPHILUS

been brackish water. The bottom was bedrock limestone. The water
was dark and extremely turbid and filled with debris. Both along the
edges and in the middle, there was considerable emergent vegetation.
The second locality is given as Touleum, Quintana Roo. This
probably is the archeological site of Tulum on the east coast.

MATERIAL EXAMINED

Holotype male, allotype female, and five male and five female paratypes
with the following locality data are in the United States National Museum:
5 miles south of Progreso, Yucatan, xi.24.63, J. R. Zimmerman. Two para-
types of each sex with same data are also in the California Academy of Sci-
ences, San Francisco, and the Departmento de Entomologia, Laboratorio de
Sanidad Vegetal, Coyoacan, Mexico. A male paratype from Touleum, Quin-
tana Roo, vili.12.49, C. J. Goodnight, is also in the University of Michigan
Museum of Zoology, Ann Arbor.

MEXICO. — YUCATAN. Progreso, 5 m. S., 36 4, 55 2, xi.24.63, JRZ
(NMSU).

Laccophilus sonorensis, new species (Fig. 308)

DIAGNOSIS. — A comparison of elytral patterns will permit separation of
sonorensis from all other species. It most resembles the sympatric L. q. quadri-
lineatus, but the pattern is almost a ‘“‘negative” of that form. Anatomically, the
two are extremely similar and, obviously, closely related. The postcoxal proc-
esses of quadrilineatus appear to project slightly more posteriorly than they do
in sonorensis, however. Megascopically, sonorensis might be mistaken for the
more common L. fasciatus terminalis, but a microscopic examination will readily
show the differences. L. vacaensis thermophilus occurs in the same area and,
like sonorensis, lacks a metacoxal file in the male and has widespread teeth on
the ovipositor; but it is an irrorated species with a different elytral pattern and
is generally much smaller. No other sympatric Laccophilus species would be
confused with sonorensis.

DESCRIPTION. — Medium to large (length, 4.5 to 6.0 mm; width, 2.5 to
3.2 mm), yellow and brown, non-irrorated species; metacoxal file absent; pro-
sternal process short; ovipositor more rakelike than sawlike. COLOR. Head
and pronotum: pale yellow, faded near margins. Elytra: yellow with brown
markings; elytral suture dark reddish-brown; epipleura pale anteriorly and
sometimes darkened to translucent reddish-brown posteriorly. Tergite VII:
brown at base, but posterior half yellow. Venter and genitalia: pale yellow
tinged with reddish-brown. ANATOMY. Microreticulation: weakly double on
head and pronotum with inner cellules nearly as distinct as secondary network;
elytra single and more pebbled, especially females. Head: supraclypeal seam
deeply impressed and closely parallel to the margin. Pronotum: mid-discal de-
pression longitudinally elongate; WH/PW, 0.69; LP/PW, 0.42; posterior mar-

JAMES R. ZIMMERMAN 189

WY VILLI =
a, MMS TEE jer pA
5.6 4

q. tehuanensis

5.0 5.2 5.4 5.6 5.8 6.0 6.2 6.4

N UM BERS

5.0 5.2 5.4 5.6 5.8 6.0 6.2 6.4

TOM VAG ESE NG eh cH (mim)

Figure 20. Histograms of length in Laccophilus quadrilineatus and L.
sonorensis. Males are shown crosshatched; females, stippled.

gin sharply deflected forming a more acute apical angle than is common in the
genus. Elytra: apices somewhat truncated; epipleura of females with anterior
wide portion extending and widening posteriorly as far as the third abdominal
segment instead of constricting at about the first as in most of the genus.
Venter: prosternal process short and broad with raised median crest; postcoxal
processes projecting only slightly beyond the midline; outline of last ventral seg-

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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JAMES R. ZIMMERMAN 191

ment subtriangular with rounded crest in females; more rounded with produced
apex in males; an asymmetrical curving crest present; lateral surfaces of last
segment in both sexes sloping rather sharply upward when viewed ventrally
causing the posterior margin to be less rounded and more angular; apex of fe-
male segment reflexed ventrally; a few rugae and numerous setigerous punctures
accumulating toward posterior margin. Legs: pro- and mesotarsi enlarged in a
dorsoventral plane and dilated slightly laterally; fifth protarsal segment at least
twice as long as corresponding fourth and fifth mesotarsal segment slightly more
than twice as long as corresponding fourth; palettes easily visible at 20 power
magnification; pro- and mesofemoral marginal setae very long (some at least as
long as femur is wide); profemoral ones (6 to 7) shorter and finer than meso-
femoral ones (7 or 8). Genitalia: oval plate with elongated tip; its ventral
crest distinct only near apex; numerous fine raised lines on either side of crest;
aedeagus wider in distal than in basal half, but narrowing near apex to a small
slightly bent and rounded tip; parameres of nearly equal length; right one with
elongated apex; ovipositor with about 11 pair of widely spaced, but triangular
teeth.

NOMENCLATURAL NOTES. — It is obvious that sonorensis is
closely related to L. q. quadrilineatus. Anatomically, they are virtu-
ally identical; but the elytral patterns and size are different (Table 16,
17). L. sonorensis males are about 0.45 mm smaller than quadri-
lineatus males, and the females are 0.3 mm smaller than the latter
(fig. 20). In the absence of intergrades and with a considerable area
of overlap, it seems necessary at the present time to consider the two
as distinct.

VARIATION. — Males are smaller than females, but the smallest
individual from a sample of about 85 specimens was a female which
measured only 4.97 mm long. The next smallest individuals were
two males that measured 5.18 mm. Also, the largest male was as
large as the largest female (about 6.05 mm long). The population
from La Paz, Baja California, may be larger than those from Sonora,
Arizona, and New Mexico; but the samples are too small to give con-
clusive results (Table 17). There seems to be no difference between
the elytral patterns of males and females. The elytra have a discal
blotch that has a variable amount of clearing in the center and is ex-
tended anteriorly in fingerlike projections similar to that of quadri-
lineatus. The degree to which the projections are joined to the blotch
varies from extensive to almost none.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—The northern limit of this species is confined to southern New

MEM. AMER. ENT. SOC., 26

192 THE GENUS LACCOPHILUS

Mexico and Arizona (fig. 19). The easternmost records are in Dona
Ana County, New Mexico, and Samalayuca, Chihuahua, which lies
about 30 miles south of El Paso, Texas. The northwestern record is
Mono County, California; and in Sonora, it has been taken about as
far south as Hermosillo. It has been collected on two occasions near
La Paz, Baja California, but nothing is known about its distribution
farther north on the peninsula. The gap in southwestern Arizona is
due to a lack of good collections from that area.

This species, which occurs in the same ponds as quadrilineatus,
seems to prefer temporary situations in the flat areas between the iso-
lated ranges in the southwestern United States and northern Mexico.
It is a desert species that can, apparently, tolerate the high water tem-
peratures that must frequently occur. It must be highly mobile and
readily fly from one drying puddle to another which may be 20 or
more miles distant.

MATERIAL EXAMINED

Holotype male, allotype female, and five paratypes with the following data
are in the California Academy of Sciences, San Francisco: La Paz, Baja Cali-
fornia Sur, vili.28.60, A. G. Smith. Two more males and one female paratypes
(8.2 m. W. of La Paz, Baja California Sur, xii.31.58, H. B. Leech) are also in
the California Academy of Sciences. A male and female paratype from 5 miles
south of Animas, Hidalgo County, vii.24.61, J. R. Zimmerman, are in each of
the following: the United States National Museum, the University of Michigan
Museum of Zoology, Ann Arbor, and the Departmento de Entomologia, Labora-
torio de Vegetal Sanidad, Coyoacan, D. F., Mexico.

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Hua-
chucha City, 2 6, 1 2, ix.6.61 (NMSU). Tombstone, 5 m. N., | 2, ix.6.61;
Rodeo (N.M.), 2 m. NW., 4 6, 4 2, 1x.5.61, JRZ (NMSU). Maricopa
County. Gillespie Dam, 1 9, vili.18.64, Arnett & Van Tassel (CUA). Pima
County. Tucson, 1 ¢, ix—.33, O. Bryant (CAS). Pinal County. Superior,
1 6, vii.15.27, E. G. Graham (CNL). Picacho Pass, 1 2, ix.12.54, J. C. Hall
(DAV). Santa Cruz County. Pena Blanca, 1 ¢, viii.9.61; 1 2, viii.5.62,
Arnett & Van Tassel (CUA). CALIFORNIA. Mono County. Mammoth,
2 4, vili.17.52, P. S. Bartholomew (CAS). Riverside County. Blythe, 1 2,
vi.7.62, R. M. Hardman (BERK). NEW MEXICO. Dona Ana County.
Jornada Range, Taylor Wells, 1 4, vii.20.63; 3 6, vii.28.63, K. L. McWilliams
(NMSU). Hidalgo County. Animas, 5 m. S., 14 4, 12 9, vii.24.61, JRZ
(NMSU); 1m. S., 1 4, 1 2, vii.31.65, H. B. Leech (CAS). Gary, 5 m. W.,
8 6, 7 2, vii.24.61; Roadforks, 4 6, 2 2, ix.5.61, JRZ (NMSU). Double
Adobe Ranch, Animas Mtns., 2 9, viii.15.52, H. B. Leech (CAS).

JAMES R. ZIMMERMAN 193

LACCOPHILUS BIGUTTATUS e

LACCOPHILUS UNDATUS °

LACCOPHILUS SCHWARZI s

Figure 21. Distribution of Laccophilus biguttatus, L. undatus, and L.
schwarzi.

MEXICO. — BAJA CALIFORNIA SUR. La Paz, 7 ¢, 2 9, viii.28.60, A. G.
Smith; 8.2 m. W., 2 6, 1 2, xii.31.58, H. B. Leech (CAS). CHIHUAHUA.
Samalayuca, 1 92, viii.6.50, R. F. Smith (AMNH). SONORA. Hermosillo,
1 @, viii.15-20.53; 50 km. W., Rancho San Francisco, 3 9, viii.2.53, B. Malkin;
San Carlos Bay, 1 4, viii.10.60, Arnaud, Ross, & Rentz; Rancho Montijo, 25 m.
E. Puerto Kino, 1 @, viii.1.50, J. Figg-Hoblyn (CAS).

Laccophilus biguttatus Kirby (Figs. 201-208, 321)

Laccophilus biguttatus Kirby, 1837, p. 69. Type: British Museum (Natural
History), Boreal America; Leng, 1920, p. 77; Zimmermann, 1920, p. 25;
Balfour-Browne, J., 1944, p. 345; Blackwelder and Blackwelder, 1948, p. 3.

Laccophilus inconspicuus Fall, 1917, p. 164; Leng, 1920, p. 76; Zimmermann,
19205 ps2):

DIAGNOSIS. — This is the only Laccophilus occurring in North America
with a pale unicolorous or concolorous elytra. All others are irrorated, varie-
gated, marmorated or spotted, or much darker. It resembles L. mexicanus, an
irrorated species, but lacks a file in the male. L. biguttatus also his thickened,
darkened antennal segments, a character possessed as well by the strongly pat-
terned undatus.

DESCRIPTION. — Medium (length, 3.9 to 4.8 mm; width, 2.3 to 2.6 mm),
concolorous, brownish-yellow, non-irrorated species; metacoxal file absent, pro-
sternal process short; ovipositor sawlike. COLOR. Head: generally brownish-
yellow above and beneath, but darker on occiput, brownish on tips of maxillary
and labial palpi and on the distal halves of the last seven antennal segments.

MEM. AMER. ENT. SOC., 26

194 THE GENUS LACCOPHILUS

Pronotum: about the same as head and with a darker area on the anterior mar-
gin corresponding to the one on the head. Elytra: generally unicolorous light
brownish-yellow to light brown; some suggestion of pattern along the anterior
half of the lateral margins, subbasally, and just behind the middle; epipleura
entirely pale. Tergite VIII: light to medium brown. Venter: variable from
entirely brownish-yellow to nearly reddish-black or black; legs usually remain
pale yellowish-brown tinged with red. Genitalia: variable in the same fashion
as the venter. ANATOMY. Microreticulation: faintly double on the head and
pronotum, but individual cellules strongly impressed; single on the elytra; sur-
face shining. Head: antennal joints thicker than other North American Lac-
cophilus; supraclypeal seam parallel, but separated somewhat from the margin.
Pronotum: WH/PW, 0.68; LP/PW, 0.40. Elytra: apices slightly truncated;
female epipleura without flange. Venter: prosternal process with well-defined
crest from base to apex; postcoxal processes rounded and laterally projecting
well posterior to midline; last ventral abdominal segments in both sexes neither
produced nor truncated; median crest on female segment marked and rooflike
at apex; male crest nearly symmetrical so that the outline is similar in the two
sexes; scattered setigerous punctures. Legs: male pro- and mesotarsi enlarged
slightly in a dorsoventral plane; palettes easily visible at 20 power magnifica-
tion; fifth tarsal segment on both pair of front legs about one and three-fifths
as long as corresponding fourth; setae on profemoral margin (7 or 8) similar
in diameter, length and number to mesofemoral ones. Genitalia: oval plate
produced to point; its median crest with leftward curvature; somewhat stronger
than the raised lines which are present on either side; aedeagus strongly curved
at about half its length, expanding before the apex, but narrowing near tip;
setae projecting from apex of right paramere instead of subapically as in most
members of the genus; ovipositor with about 15 sawlike teeth on each row.

NOMENCLATURAL NOTES. — The biguttatus of Kirby (1837)
was long unrecognized and was ascribed to fasciatus or proximus.
Sharp’s (1882a) monograph does not include the species. Fall’s in-
conspicuus was recognized as a synonym by J. Balfour-Browne (1944) ;
and, at the same time, its identity was clarified. Kirby’s description
noted the darkened antennal segments and palpi; but his references to
spots (that are weak at best and, perhaps, partly due to translucence )
delayed the recognition of this distinctive species.

L. biguttatus is strongly similar to L. minutus (L.) of the Old
World, as noted by Kirby, and, eventually, may prove to be a race of
an extremely widespread polytypic species. The genitalia, thickened
and darkened antennae, lack of file, microreticulation, size, and shape
are all close; and only the elytral pattern differs. The last is a fea-
ture that varies considerably in other races of several North American
species.

195

ZIMMERMAN

JAMES R.

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MEM. AMER. ENT. SOC., 26

196 THE GENUS LACCOPHILUS

VARIATION. — The venter varies from pale yellowish-brown to
nearly all black in some specimens. This feature is not clearly geo-
graphic, but is generally uniform for any one sample.

Females appear to be slightly larger than males, but with a large
overlap (Table 18). The peak in the frequency distribution of length
for females is about 4.4 mm, and that for males is about 4.25 mm
(fig. 22). The largest females are slightly larger than the largest
males, and the same relationship holds for the smallest of each sex as
well. In samples from six different areas females always had larger
values than the males from the same locality.

Quantitative comparisons of seven samples from New Hampshire
to the Yukon give no indication of any racial differentiation of bi-
guttatus within North America. Differences appear to be more local

biguttatus

NUMBERS

TOTAL LENGTH (mm)

Figure 22. Histograms of length in Laccophilus biguttatus. Males shown
in crosshatch; females, stippled.

than regional. The three largest and most reliable samples (from
Manitoba, Alberta, and from Colorado) are similar. Males from the
Yukon (n=9) do appear to be smaller than all other males, except
those from New Hampshire (n= 3), and may reflect the extreme en-
vironment of the northern climate.

JAMES R. ZIMMERMAN 197

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This is the only boreal species of Laccophilus in North America.
It occurs in all the northern tier of states and as far south as north-
western New Mexico and the Sierra Nevadas of California. It is known
from all provinces from Quebec west. It is not known how far north
the range extends in eastern Canada; but it does occur in the North-
west Territories, the Yukon, and Alaska (fig. 21).

L. biguttatus seems to be the commonest Laccophilus in the in-
terior provinces. I have collected this species only in the Chuska
Mountains of northwestern New Mexico and at an altitude of about
9000 feet. It was in mountain meadow ponds that were choked with
water weed and algae, but with clear water. Fall quotes Sherman
that, in New Hampshire, they are taken at higher altitudes of the
White Mountains. In North Dakota, Wisconsin and Michigan, they
had to be taken at much lower altitudes than the two previously cited
localities.

MATERIAL EXAMINED

CANADA. — ALBERTA. Calgary, Calgary, 1, x.23.27, O. Bryant (CAS).
Cypress Hills, 1, viii.25.27 (USNM); 1 @2, vi.13.24 (DAV). Edmonton, 2
paratypes (inconspicuus Fall); 4, iv.12.17; 14, iv.22.17, F. S. Carr (USNM);
19, iv.4.18, F. S. Carr; 30, viii.19.27, O. Bryant; 4, vi.12.17, F. S. Carr (CAS).
2 B52 5 VDI 3 SG, MONS DA Se wills 1 G5 wre Il Gy il Qe my 7cIe
F. S. Carr (MCZ). High River, 1, vi.25.27, O. Bryant (CAS). Medicine Hat,
1 4, viii.21.23, F. S. Carr (FM); 1, vii.29.—, H. J. Rayner (CAS). Red Deer,
1, vii.19— (USNM). Tofield, 1, x.24.24; Turner Valley, 25, v.24.29, O. Bry-
ant (CAS). Waghorn, 10 (USNM). Wastoc, 1 6, x.12.21, F. S. Carr (DAV).
BRITISH COLUMBIA. Clinton, 2, vi.18.31, R. Hopping; 2, x.11.43, H. B.
Leech & C. V. Morgan; Copper Mtn., 1, v.15.43, G. S. Smith; Enderly, 1,
iv.1.45, H. B. Leech; Fernie, 11, ix.16.28, O. Bryant; Lumby, 1, v.11.33, E. R.
Backell (CAS). Lac Le Hache, 3 4, viii.—.59, J. Robertson (LACM). Nicole
Lake, 1 4, viii.25.32, A. E. Thrupp (BERK); 3, viii.25.32, A. Thrupp; Oliver,
iv.25.33, E. R. Backell (CAS). Paxton Valley, 1 2, vi.22.23, A. E. Thrupp
(BERK); 6, vii.22.53, A. E. Thrupp (CAS); 1 6, vi.22.23 (LACM). Similka-
meen, Taylor Swamp, 2, iii.29.41, High (CNL); Similkameen, 2, i1i.29.41,
H. B. Leech; Upper Hat Ck., 1, viii.25.32; 1, vili.29.33, G. J. Spencer (CAS).
Vernon, 2 6, 5 2, ix.2.40, H. B. Leech (LACM). DISTRICT OF MAC-
KENZIE. Aklavik, 3, viii.5.30; 2, ix.5.30; Hay River, 3, ix.30.29; 16, vii.12.30,
O. Bryant (CAS). MANITOBA. Aweme, 7, vi.16.04, Criddle; Piquitenay, 2,
vii.24.17, J. B. Wallis (USNM). Stonewall, 1, v.11.24, J. B. Wallis (CNL).
Treesbank, 11 ¢, 8 92, vii.10.25, J. B. Wallis (AMNH); 8 6, 12 9, vii.10.25
(MCZ). Winnipeg, 6, viii.16.16; 2, v.27.24, J. B. Wallis (USNM); 1, v.4.24,

MEM. AMER. ENT. SOC., 26

198 THE GENUS LACCOPHILUS

J. B. Wallis (CNL); 1 6, ix.2.16, J. B. Wallis (CNG); 1 2, v.1.—; 1 6, 1 2,
v.8.—, J. B. Wallis (AMNH); 1 6, viii.16.16, J. B. Wallis (CAS). ONTARIO.
Nipigon, 1 2, viii.29.27, Darlington (MCZ). Sioux Lookout, 1 6, 1 2, ix.5.16
(CNG). QUEBEC. Abitibi Post, 5, vii.23.35; Duparquet, 4, viii.2.35; 3,
v.15.38; 5, viii.28.39; 5, vii.21.43; 1, ix.2.43, G. Stace Smith (CAS). Montreal,
3 (USNM); 2 6, 2 2, vii——, Chagnon (AMNH). Notoshquan, 1, xi.8.29,
W. J. Brown (CAS). Strathcone, 1, v—.05 (USNM). SASKATCHEWAN.
Lost Mtn. Lake, 1, JST (CNL). Murray (Lake?), NW. of Battleford, 1,
v.28.39 (CNL). Redberry Lake, 2, vi.24.39; Redvers, 2 (CNL); 1 ¢ (MCZ).
Regina, 1, v.30.38, C. Shaw (CAS); 1 ¢, iv.15.44, C. C. Shaw (MCZ). Stur-
geon (Lake?), 2, vi.24.39 (CNL). Yorkton, 14, vi.5.47, C. C. Shaw (CAS);
1 &, iv.24.45, C. C. Shaw (MCZ). YUKON. Mayo, 6 6,5 9, vii.3.55, F. S.
Carr (CARR).

UNITED STATES OF AMERICA. — ALASKA. Fairbanks, 2 2, viii.4.58,
Lindroth (CAS); Fairbanks, Farmers’ Loop, 2 2, viii.11.57; Glenn Hwy., Mile
Post 86, 1 4, viii.28.57; Alaska Hwy., Mile Post 1370, 1 6, viii.10.57, E. L.
Kessel (CAS). CALIFORNIA. Tulare County. Sequoia Natl. Park, 1 ¢,
F. E. Winters (CAS). COLORADO. Boulder, Grant or Larimer Counties.
Battle Mtn. Dam, 15, vii.17.37, E. B. Andrews (CAS). Rocky M.N.P., Glacier
Basin, 2, vii.—.37, E. B. Andrews (CNL); 2, vii.1.37 (USNM); 3 @, vii.1.37,
H. B. Leech (FM); 2 ¢, vii.1.37, J. G. Thorndike (AMNH). R.M.N.P.,
8500 feet, 1, F. E. Winters (CNL); Storm Pass, 3, vii.8.38, E. B. Andrews
(CAS). Glacier Basin, 2 ¢, viii.1.37, Andrews (MCZ). Rocky M.N.P.,
11 6, 10 2, F. E. Winters (CAS). Jackson County. Cameron Pass, 15,
vil.19.41, H. P. Chandler (CAS). Muddy Pass Jct., Hwy. 40, 2 6, 2 9,
vii.3.63, R. D. Anderson (RDA). Pinegrass Pk., 1, viii.8.25, Statterthwait
(USNM). Park County. Kenosha Pass, 10,000 feet, 3, vii.16.38, J. W. Green
(CAS). MICHIGAN. Cheyboygan County. Smith’s Bog, 1 é, vii.5.52, P. J.
Spangler (USNM). Livingston County. E. S. George Reserve, 7, vi.i0.50,
F. N. Young (CAS). Marquette County. Marquette, 7, vii.1-2.16 (USNM);
1 Q, vii.2.16, J. D. Sherman, Jr.. (AMNH). MONTANA. Carbon County.
E. Rosebud L., 1 2, vi.19.61 (DAV). NEW YORK. Essex County. Mt.
Marcy, L. Tear, 1 (USNM). Tompkins County. Ithaca, 4, vi.9.17; 2, vi.17.17;
2, v.21.26, H. Dietrich (CNL). NEW HAMPSHIRE. Coos County. Mt.
Washington, 1 (USNM). Peabody Spring, 5000 feet, 1 2, ix.—.—, J. Sher-
man, Jr. (AMNH). Starr L., 4800 feet, 1 (USNM). Storm L., 4930 feet,
1 (USNM). Crafton County. Rumney, 1 ¢, 2 2, iv.21.16; 1 6, 1 2, iv.22.16;
1 6, 1v.17.27, Darlington (MCZ). NEW MEXICO. San Juan County. Chuska
Mtns., Washington Pass, 9000 feet, 50+, x.5.68, JRZ (NMSU). NORTH DA-
KOTA. Benson County. Ft. Totten Lake, 8, vi.21.21, C. K. Sibley; 14, viii.22.20
(CNL). Bottineau County. Turtle Mtns., Fish Lake, 8, vii.13.21; L. Meti-
gishe, 1, vii.22.21; Dion Lake, vii.15.21, C. K. Sibley (CNL). Ramsey County.
Devil’s Lake, S. Odessa Pond, 1, vii.5.21; N. Odessa Pond, 1, viii.S.21, C. K.
Sibley (CNL). VERMONT. Chittenden County. Mt. Mansfield, Lake Cloud,

JAMES R. ZIMMERMAN 199

—™.
Ppp >

Ze
LH LACCOPHILUS G. GENTILIS 4
{ LACCOPHILUS G. SUAVIS 4

LACCOPHILUS HORNI ©
LACCOPHILUS LEECHI °

LACCOPHILUS YOUNGI ®

Figure. 23. Distribution of Laccophilus gentilis, L. duplex, L. horni, L.
leechi, and L. youngi.

1, iv.19.13 (USNM). WISCONSIN. Oneida County. Three Lakes, 11, 1,
ix.—.04 (USNM); 3 6, 1 2, W. S. Marshall (AMNH). WYOMING. Yellow-
stone Natl. Park, 8000 feet, 1 2, vili.17.33, E. B. Andrews (CAS).

Laccophilus horni Van den Branden (Figs. 209-216, 324)

Laccophilus lateralis Horn, 1883, p. 282. Lectotype, male, Academy of Natu-
tal Sciences, Philadelphia, No. 2961, Arizona.

Laccophilus horni Van den Branden, 1885, p. 21; Leng, 1920, p. 76; Zimmer-
mann, 1920, p. 19.

DIAGNOSIS. — Elytral pattern, pronotal markings, and absence of meta-
coxal file separate this species from all other ones in North America. Only
pictus and leechi have an elytral pattern with yellow spots on a black or nearly
black background. L. pictus has a file and darkened epipleura. The epipleura
are pale in horni, and the arrangement of the spots is different (figs. 300-304,
324). L. horni and L. leechi have very similar elytral patterns and lack files,
but the pronotal markings of /eechi are more extensive. Also, horni has a mean
length of about 4.5 mm; and Jeechi is only about 3.5 mm long.

MEM. AMER. ENT. SOC.. 26

200 THE GENUS LACCOPHILUS

DESCRIPTION. — Medium (length, 4.10 to 4.95 mm; width, 2.45 to 3.10
mm), dark brown or black species with yellow spots; metacoxal file absent,
prosternal process short; sawlike ovipositor. COLOR. Head: pale brownish-
yellow faintly tinged with red except for dark reddish-brown occiput between
the eyes; appendages pale. Pronotum: pale brownish-yellow tinged with red in
center and laterally, but with dark reddish-brown bilobed spots on front between
the eyes and at the apex; entire posterior margin translucent brownish-red.
Elytra: strongly marked spotted pattern of yellow and very dark reddish-brown
or black; epipleura pale anteriorly, posterior half may darken to reddish-brown.
Tergite VIII: pale yellow-brown. Venter: medium yellowish-brown; pro- and
mesolegs generally paler, darker at the bases of mesocoxae and on the meta-
coxal plates around the base of the hind legs; lateral margins of plates also dark
reddish-brown. Genitalia: variably pale reddish yellow-brown to reddish-brown.
ANATOMY. Microreticulation: double on head, pronotum, and elytra; surface
shining. Head: supraclypeal seam arching slightly above the clypeal margin.
Pronotum: relatively short since angle between it and the elytra approaches 180
degrees; WH/PW, 0.69; LP/PW, 0.38; surface roughened, but shiny. Elytra:
apical truncation barely perceptible; females without epipleural flange. Venter:
prosternal process short, slender, with well-defined crest; postcoxal processes
not laterally projecting past the midline, but instead forming a nearly straight
line or bending slightly more anteriorly than the midline; last abdominal seg-
ments of both sexes not truncated, but perhaps somewhat elongated to subtri-
angular; deep, marginal groove extending almost the entire lateral posterior
margin to the crest in females and about half the length of the margin on either
side of the apex in males; median crest nearly symmetrical in both sexes; numer-
ous setigerous punctures scattered over the surface, thickest on crest. Legs:
male pro- and mesotarsi enlarged slightly in a dorsoventral plane; fifth tarsal
segments on both pair of front legs one and three-quarters to twice as long as
corresponding fourth; mesofemoral setae (8 to 9) much larger than profemoral
ones (7); metatibial spur nearly as long as two tarsal segments. Genitalia: oval
plate with produced acuminate point; its median crest only distinctly raised near
apex, but apparent anteriorly with a bend to the right; a few weak raised lines
on either side; aedeagus simple and tapering gradually along entire length; right
paramere with blunt apex; ovipositor with about 14 sawlike teeth.

NOMENCLATURAL NOTES. — Horn (1883) used a preoccu-
pied name in describing this species, and this was corrected by Van
den Branden (1884).

VARIATION. — Although the yellow elytral spots vary in size,
there is no particular geographic trend. The subbasal spots may be
completely joined or separated into two or three discreet ones.

Females tend to be slightly larger than males. The peak fre-
quency for females lies at about 4.6 mm and that for males at about
4.5 mm (fig. 25). Ina total of 254 specimens measured, however,

201

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

202 THE GENUS LACCOPHILUS

undatus

N U M 6 E RS

gentilis gentilis

3.2 3.4 3.6 3.8 4.0 4.2
TOTAL LENGTH (mm)

Figure 24. Histograms of length in Laccophilus gentilis, L. ovatus, L.
duplex, and L. undatus. Males shown in crosshatch; females, stippled.

JAMES R. ZIMMERMAN 203

the largest individual was a male that was about 5.0 mm long. There
may be some geographical variation in the size of females compared
to males. In two samples (Table 19) from Arizona and Sonora,
there is a fairly good separation of the means of females and males.
The mean length of females over males in the Arizona sample is highly
significant, but from Jalisco south, there is no difference. Sonoran
region populations are longer, but with the large gap in samples, it
cannot be determined if the difference is racial, clinal, or local.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— The distribution as now known of Laccophilus horni is separated
into four areas (fig. 23). One is in southeastern Arizona and north-
eastern Sonora, another in the Davis Mountains in Texas, the third in
San Luis Potosi, and the last in a fairly continuous chain from Na-
yarit to Oaxaca. Some of the separation appears to be due to a lack
of collecting. It seems likely that there should be populations almost
continuously from the one in northeastern Sonora to Nayarit. On the
other hand, the Davis Mountain population may be truly allopatric to
any other populations. There is a little suitable habitat between it
and populations to the west or south. The San Luis Potosi localities
are close enough to those in Michoacan to suspect that they are not
truly allopatric populations.

L. horni has a high fidelity for stream pools occurring in oak
woodland. In all parts of its range it is found near oak in hilly terrain.
It can be collected anywhere between 1000 and 5000 feet in streams
or pools that have granitic gravelly bottom. It prefers to stay in ex-
tremely shallow water (3 inches or less), darting from one pebble to
the next. The easiest way to collect this species is to strain some
gravel with the rim of a metal sieve just above the water surface and
allow the specimens to swim upward where they can be picked up with
a forceps. They are probably one of the more difficult species to col-
lect in large numbers, which probably explains why they are seldom
abundant in collections.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. Cochise County. Chiraca-
hua Mtns., 1, v.9.—, Hubbard & Schwarz (USNM); H. Martyr L., 2 6, 2 2,
ix.5.61, JRZ (NMSU); Portal, 1 ¢, vilim.58, P. Opler (BERK); SWRS, 1,
vil.4.56, O. L. Cartwright (USNM); 1 6, 1 2, vii.22.56, E. Ordway; 1 °,
vii.19.57, M. Stathen (AMNH); Cave Ck. Cn., 1 2, ix.13-14.—, B. Malkin

MEM. AMER. ENT. SOC., 26

204 THE GENUS LACCOPHILUS

4.2 4.4 4.6 4.8 5.0

TOTAL LENGTH (mm)

Figure 25. Histograms of length in Laccophilus horni. Males shown in
crosshatch; females, stippled.

(FM). Huachuca Mtns., Bear Canyon, 1, v.8.53, A. & H. Dietrich (CNL).
Gila County. Pinal Mtns., 1, iv—.—, D. K. Duncan (CNL); 1, vi.15.47,
F. H. Parker (CAS). Graham County. Graham Mtn., Marijilda Canyon,
11 é, 16 9, ix.12.52, B. Malkin & V. E. Thatcher (FM). Pima County.
Baboquivari, 2, —.—.24, C. C. Poling (CNL); 7, ix.24.33, Bryant (CAS).
Catalina Mtns., 2, iv.18-v.16.—, Hubbard & Schwarz (USNM); Sabino Can-
yon, 2, vii.11.49, D. J. & J. N. Knull (USNM); 1 2, 1x.6.63, V. L. Vesterby
(DAV); 25, xii.28.50, R. S. Beal (BERK); Molino Basin, 3, vi.11.58 (CAS);
1 6, vi.9.54, R. S. Beal (NMSU); 3 6, 2 9, vi.9.54, R. S. Beal (BERK); 1 2,
xi.27.27, F. H. Parker; 1 9, x.16.43, J. Hendrickson (ARI). Santa Cruz
County. Nogales, 2, iv.—.97, Koebele (CAS). Pena Blanca, Pajarito Mtns.,
4 4,3 Q, vii.26.64, R. H. Arnett & E. Van Tassel (CUA). Santa Rita Moun-
tains, 1 2, xi.6.24, A. A. Nichol (MCHS); Madera Canyon, 1 2, x.2.63, V. L.
Vesterby (DAV); 4, vii.26.55, F. X. Williams (CAS); 1 2, vi.6.56, A. Menke
(LACM); Florida Canyon, 3 2, vili.6.62; 1 9, i1x.7.61; 2 6, 1 92, xii.2.61,
JRZ (NMSU). Tumacacori Mtns., 12 6, 6 2, vil.27.65, H. B. Leech (CAS).
TEXAS. Davis County. Davis Mtns., Madero Canyon, 1, vi.14.16, F. M.
Gaige (USNM); Phantom L., 2, viii.20.16, F. M. Gaige (CAS). County, 4 é,
4 9, vili.10.16 (MCHS).

MEXICO. — JALISCO. Mazamitla, 17 m. S., 1 2, xii.5.48, H. B. Leech
(CAS). Between Autlan and Union de Tula, R. San Pedro, 3 6, 7 9, iii.28.64,

JAMES R. ZIMMERMAN 205

JRZ (NMSU). MEXICO. Tonatico, 5 6, 2 2, viii.29.62, JRZ (NMSU).
MICHOACAN. Tzitzio, 3 2, vil.27.62; 3 m. S., 17 6, 15 @, viii.22.63, JRZ
(NMSU). Uruapan, 10m.S., 1 4,1 9, i1i.26.64, JRZ (NMSU). MORELOS.
Cuerna (vaca?), 1 29, v—.—, Barrett (CAS). NAYARIT. Sierra de Zapotan,
2 2, xi.—.42, Eugenio Paredes (CAS). Tepec, 25 m. SE., 4 ¢, 2 9, xi.23.48,
H. B. Leech; 25 km. S., 1 6, 1 2, ix.24.53, B. Malkin (CAS). OAXACA.
El Tule, 1 6, ix.6.64; Huajuapan, 2 km. S., 4 6, 4 9, 1x.4.64; 17 km. NE.,
1 2, ix.3.64; Oaxaca, 3 m. S., 1 92, viii.26.63, JRZ (NMSU). SAN LUIS
POTOSI. Cuidad del Maiz, 14-15 m. E., 1 6, 2 2, ix.19.48; San Luis Potosi,
2m. S., 2 6, xi.21.48, H. B. Leech (CAS). SONORA. MHuachineura, 11 4,
7 Q, vii.25.63, JRZ (NMSU).

Laccophilus leechi, new species (Figs. 217-224, 325)

DIAGNOSIS. — Small size, yellow spotted elytral pattern, darkened prono-
tal disc, and no metacoxal file separates Jeechi from all other North American
Laccophilus. L. leechi most resembles horni; and if it were not for their ap-
parent sympatry, it would be described as a race of the latter. It differs from
horni only in its much smaller size, in the slightly different conformation of the
aedeagus, and in the more broadly pigmented pronotum. L. leechi, L. gentilis
suavis and L. oscillator laevipennis are about the same size and sympatric, but
suavis and laevipennis have nearly complete subbasal elytral bands and much
lighter pronota. L. g. suavis also has a file. The aedeagi of it and Jeechi are
quite different, but L. oscillator laevipennis has a similar aedeagus. All other
species in western Mexico are either much larger or have irrorated patterns.

DESCRIPTION. — Small (length, 3.5 to 3.8 mm; width, 2.1 to 2.2 mm),
dark reddish-brown to black species with yellow spots; metacoxal file absent;
prosternal process intermediate; ovipositor sawlike. COLOR. Head: pale yel-
low above and beneath except dark reddish-brown on occiput between the eyes;
appendages except mandibles, pale yellow. Pronotum: large reddish-brown
blotch extending from between the eyes to the apex; remainder pale yellowish-
brown. Elytra: strongly contrasting, spotted pattern of yellow and dark reddish-
brown or black; epipleura entirely pale or with posterior portion of translucent
reddish-brown. Tergite VIII: distal half pale yellowish-brown, basal half dark
brown. Venter: generally pale yellowish-brown tinged with red except for
around mesocoxal bases and lateral margin of the metacoxal plates, or entire
metasternum and venter strongly suffused with black or dark reddish-brown;
pro- and mesolegs generally pale brownish-yellow; hind legs darker and tinged
with red. Genitalia: variable from pale brownish-yellow to reddish-brown.
ANATOMY. Microreticulation: secondary reticulation strongly developed and
individual cellules only weakly visible on head; pronotum and elytra double
and with inner cellules showing distinctly only on the elytra; surface shining.
Head: supraclypeal seam closely parallel to margin. Pronotum: relatively short;
WH/PW, 0.69; LP/PW, 0.38. Elytra: apical truncation barely perceptible;
female epipleura without flange (only six females examined). Venter: pro-
sternal process intermediate, extending nearly as far posteriorly as an imaginary

MEM. AMER. ENT. SOC., 26

206 THE GENUS LACCOPHILUS

line drawn across the rear of the mesocoxal cavities and the groove which re-
ceives it extending beyond that line; postcoxal processes not laterally extending
beyond the midline, but with hind margins extending straight across or curving
slightly anteriorly; last segment of male and female similar, not truncated, but
rounded or subtriangular with nearly symmetrical median crests; scattered setig-
erous punctures and a marginate groove on female segment. Legs: mesotarsi
of males enlarged slightly at most; fifth tarsal segment of both pair of front legs
about twice as long as corresponding fourth; palettes barely visible at 20 power
magnification; profemoral setae (5 or 6) occurring in two distinct clumps and
as long as but finer than those on mesofemur (7 to 8). Genitalia: oval plate
with produced, acuminate tip and median crest; aedeagus curving only slightly
over posterior three-quarters of length, abruptly narrowed near tip; ovipositor
with about 18 pair of sawlike teeth.

VARIATION. — The most obvious color variation is a dark ven-
ter on the single specimen taken in Jalisco. Dark pigment suffuses
most of the metacoxal plates and abdomen. Those from Sinaloa had
dark pigment confined to leg bases and along the metacoxal margin.
The effect is striking enough to suggest that perhaps two different
races may exist, but not enough is known yet to determine this.

Males and females appear to be about the same size; but with only
nine specimens, little can be said about any difference (Table 19).

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This species has been collected at only two localities and on but
three occasions (fig. 23). One locality is at the base of the Sierra
Madre Occidental eight miles east of Concordia on the highway be-
tween Mazatlan, Sinaloa, and Durango. L. leechi was taken there on
two different occasions. The stream bed has many quiet pools and
large boulders along its course. The insects seem to be both among
the rocks and pebbles and in places where there is some mud on the
bottom. The only other place it has been collected is in the valley in
Jalisco in which the village, La Huerta, is located. In this instance a
single specimen was taken along the margin of a small stream (three
to four feet wide), which had a bottom of mud and volcanic pebbles,
grass along the margins, and no large boulders.

MATERIAL EXAMINED

Holotype male, 8 miles E. of Concordia, iv.13.68, B. Reilly and J. R. Zim-
merman; allotype female, from same locality, but taken by J. R. Zimmerman,
on xii.12.62, are both in the United States National Museum. Also one male
and one female paratype with the same data as the allotype are in the California

207

ZIMMERMAN

JAMES R.

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MEM. AMER. ENT. SOC., 26

208 THE GENUS LACCOPHILUS

Academy of Sciences, San Francisco; one female paratype, same data is in the
Departmento de Entomologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F.,
Mexico, and the University of Michigan Museum of Zoology, Ann Arbor.

MEXICO. — JALISCO. La Huerta, 5 m. N., 1 2, x.25.66, A. H. Smith &
JRZ (NMSU). SINALOA. Concordia, 8 m. E., Rd. 15, 1 6, 1 2, xii.12.68,
JRZ (NMSU).

Laccophilus undatus Aubé (Figs. 225-233, 322)

Laccophilus undatus Aubé, 1838, p. 435. Neotype: male, United States National
Museum, Bloomington, Monroe Co., Indiana, iv.20.56, J. R. Zimmerman;
Crotch, 1873, p. 400; Blatchley, 1910, p. 210; Leng, 1920, p. 77; Zimmer-
mann, 1920, p. 27; Zimmerman, 1960, p. 144.

DIAGNOSIS. — The variegated elytral pattern and darkened palpi and an-
tennae separate undatus from all other North American Laccophilus. The only
other species with darkened antennae and palpi is biguttatus, but it has nearly
unicolorous elytra of brownish-yellow color.

DESCRIPTION. — Small to medium (length, 3.2 to 4.3 mm; width, 2.0 to
2.3 mm), brown and yellow, variegated species; metacoxal file absent; proster-
nal process short; ovipositor sawlike. COLOR. Head: yellowish-brown with
a tinge of red; slightly darker at occiput; appendages with darkened tips on the
last two or three segments of the maxillary palpi; sometimes the labial palpi,
and the last six or seven segments of the antennae. Pronotum: variable reddish-
yellow brown or their combinations; usually darker on the disc and paler on
lateral margins; a darkened area on the anterior margin between the eyes.
Elytra: complex variegated pattern superimposed on a pale brownish-yellow
background tending to form two incomplete irregular transverse yellow fasciae;
one subbasally and the other just behind the middle; epipleura pale brownish-
yellow anteriorly, but darkening to reddish-brown posteriorly. Tergite VIII:
light yellowish-brown. Venter: varying combinations of light red, yellow, and
brown; prosternum, mesosternum, pro- and mesolegs usually lighter; metacoxal
plates, abdomen, and hind legs usually darker, tending more toward reddish-
brown. Genitalia: about the same as the abdomen, but usually tending more
strongly to reddish-brown. ANATOMY. Méicroreticulation: strongly impressed
and clearly double on head, pronotum, and elytra; surface shining, especially in
males. Head: supraclypeal seam diverging slightly upward at midline. Prono-
tum: WH/PW, 0.72; LP/PW, 0.39. Elytra: female without epipleural flanges;
apices without truncation. Venter: metacoxal file absent; postcoxal processes
rounded and laterally projecting well past the midline; last ventral segment of
both sexes not truncated, but evenly rounded; median crest weak and nearly
symmetrical in males; setigerous punctures sparsely distributed; outline of fe-
male segment subtriangular with well-defined median ridge; setigerous punctures
thicker near apex. Legs: pro- and mesotarsi enlarged in a dorsoventral plane;
palettes easily visible at 20 power magnification; fifth tarsal segment on both
pair of front legs about one and one-half times as long as corresponding fourth;

JAMES R. ZIMMERMAN 209

setae on profemoral margin (6) similar in length and diameter to those on
mesofemur (6 to 7). Genitalia: oval plate with strongly produced asymmetri-
cal acuminate tip; crest extending anteriorly with slight curvature to the right;
more fine lines on right than on left side of the crest; aedeagus curved sharply
at about two-thirds its length and twisted with conformation suggestive of a
raised cobra; right paramere less broadly triangular than in maculosus; left
paramere with less rounded apex than in maculosus tending to form an angle;
Ovipositor with about 16 sawlike teeth.

NOMENCLATURAL NOTES. — Aubé’s type of undatus is ap-
parently lost. There seems to be no confusion in the identity of this
species, however.

VARIATION. — Females are slightly larger than males (Table
21, fig. 24). The individuals from Indiana may be larger than those
from New York; but the western sample may be biased, since I had
more material and probably selected specimens that were in better
condition.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L. undatus occurs from Massachusetts to the vicinity around the
southern margin of Lake Michigan and from northern New York and
southern Vermont to near Washington, D.C. It ranges west as far as
Bloomington, Indiana, and the southern edge of Michigan (fig. 21).
It seems to be restricted to the shaded pools of the northeastern de-
ciduous woodlands. In southern Indiana undatus was consistently
found only in slough ponds in the drainage of a former intermittent
stream. Those ponds were acidic with a pH of 6.6. Temperatures
seldom, if ever, exceeded 30 degrees C. The bulrush Scirpus was
abundant along the margins and was the principal emergent plant.
Ludwigia and Zannichellia, which are characteristic of temporary
aquatic situations, covered the bottom.

L. undatus approaches L. schwarzi near Washington, D. C., but
so few specimens of either have been taken from there that it is not
known whether they intergrade or not. Their anatomical features are
sufficiently different to presume they do not. L. undatus is also sym-
patric without intergrading with L. biguttatus in upstate New York.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ILLINOIS. Lake County. Lake For-
est, 1 6, 2 29, ix.2— (FM). INDIANA. Lake County. Miller, 2; Pine
(Gary), 25, ix.23.30 (USNM). Monroe County. Bloomington, 4 m. N.,
DS) B. il Sy wWsSO, 26, 2 25 wiWU aos IG, sivclMsDo, iIbay thy AVaor iil GB,

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

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JAMES R. ZIMMERMAN 211

8 2, iv.26.56; 3 9, v.11.56; 3 2, vi.7.56; 9 2, x.31.56; 10 2, xi.13.56; 8 9,
xi.28.56; 2 2, vi.18.57, JRZ (NMSU). MARYLAND. Montgomery County.
Near Plummers Island, 1, ix.12.19, J. L. Wren (USNM). MASSACHUSETTS.
Berkshire County. Egremont, 1 2, C. H. Roberts (AMNH). Essex County.
North Saugus, Howlett’s Pond, 1, viii.25.07, D. H. Clemons (USNM). Middle-
sex County. Framingham, 1, iv.10.19, C. A. Frost (CNL). Norfolk County.
Brookline, 20 (USNM); 1 (CNL). Suffolk County. Boston, 1 2 (ANSP).
Cambridge, 1 (USNM). Worcester County. Brookfield, 2, x.24.24, Chamber-
lain (CNL). Northboro, 1, i—.37, C. A. Frost (BERK). MICHIGAN.
Berrien County. Stevensville, 2 6, 1 2, vi.9.63, vii.10.63, R. B. Willson
(MCHS). NEW JERSEY. Bergen County. Fort Lee, 48, ix.22.00, J. D.
Sherman, Jr. (USNM). Ramsey, 1 2, v.17.—, C. H. Roberts (AMNH).
Camden County. County, 1 6, vi.ll.—m (AMNH). Mercer County. Trenton,
1 2, iv.23.—, E. L. Dickerson (AMNH). Middlesex County. Spotwood, 1 °,
C. H. Roberts (AMNH). Ocean County. Lakehurst, 6 6, 2 2, v.29.—, C. H.
Roberts (AMNH). Sussex County. Stanhope, L. Lackawanna, 1 9, vili.14.32
(CNL). NEW YORK. Brooklyn County. Brooklyn, Cyp. Hills, 2, v.16.10,
Shoemaker (USNM). Cattaraugus County. Alleghany, 2 2 (FM). Rich-
mond County. Staten Island, Clove Valley, 1 6, vi.3.—, W. T. Davis (MCHS);
24 (USNM); 6, iv.16.05 (USNM); 2 6, 1 9, x.11.24, L. B. Woodruff
(AMNH); 1 6, 11 6, 10 9, iv.19.—; 2 2, v.20.—, C. H. Roberts (AMNH).
Erie County. Buffalo, 1 6,2 2 (AMNH). Long Island, New York City, and
Vic. Cold Spring Harbor, 1 é, viii.10.00 (AMNH). Long Island, Forest Pk.,
4, v—.— (USNM); 12, v.7.10 (USNM). New York City, 2 2, C. H. Roberts
(AMNH). Suffolk County. Orient, 25, vi.30.43, R. Latham (CNL). River-
head, 11, iv.30.40, R. Latham (CNL). Ulster County. Esopus, 6, J. D. Sher-
man, Jr. (USNM): 2 6, 2 2, ix.1.—, C. H. Roberts (AMNH). Washington
County. Washington County, 1, J. D. Sherman, Jr. (USNM). Westchester
County. Peekskill, 3 (USNM). OHIO. Jackson County. Jackson, 1, x.20.50,
P. J. Spangler (CAS). PENNSYLVANIA. Delaware County. 3 6, 2 2,
v.18.— (ANSP). Philadelphia County. Phila-Neck, 2 (USNM). _ Philadel-
phia, Wyoming, 1, iv.29.04, G. M. Greene (USNM). RHODE ISLAND.
1 ¢, 1 2, Kemp Coll. (ANSP). VERMONT. Bennington County. Benning-
ton County, 1 (USNM): 1 2 (CNG); 3 6, 2 2, ix.16.—, 2 ¢, vii—.94; 1 °,
vii.14.—, C. H. Roberts (AMNH);12 (ANSP). VIRGINIA. Fairfax County.
Black Pond, 1, ix.21.—, Shoemaker (USNM).

Laccophilus schwarzi Fall (Figs. 234-241, 323)

Laccophilus schwarzi Fall, 1917, p. 165. Holotype: male, location unknown,
Ash Grove, Virginia, Leng, 1920, p. 77.

DIAGNOSIS .— The shining surface, elytral pattern, and entirely pale an-
tennae and palpi separate this species from the similar undatus. Larger size
and pale elytral markings separate it from g. gentilis. All other species in the
eastern United States are irrorated with files.

MEM. AMER. ENT. SOC., 26

Pie: THE GENUS LACCOPHILUS

DESCRIPTION. — Small (length, 3.9 to 4.2 mm; width, 2.1 to 2.3 mm),
brownish-red and yellow variegated species; metacoxal file absent; prosternal
process intermediate; ovipositor sawlike. COLOR. Head: pale brownish-yellow,
labrum and clypeal region immediately above sometimes slightly darker; ap-
pendages pale; little or no darkening at the base of the pronotum. Pronotum:
generally pale brownish-yellow, but with dark reddish-brown marking on the
anterior margin between the eyes and a weaker one that extends along the pos-
terior margin which tends to join the anterior one across the disc. Elytra:
reddish-brown markings on a yellow background; two nearly complete trans-
verse irregular fasciae; one subbasally and the other behind the middle; epi-
pleura pale anteriorly but with dark ventral-medial edge; posterior half dark.
Tergite VIII: pale brownish-yellow. Venter and legs: generally variable from
pale reddish-yellow-brown to light reddish-brown, but noticeably darkened on
the lateral and posterior margins of the metacoxal plates and to a lesser ex-
tent around the postcoxal cavities. Genitalia: variably reddish-yellow-brown.
ANATOMY. Microreticulation: strongly double on head, pronotum, and elytra
with individual cellule outlines weak; surface shining. Head: supraclypeal seam
continuing parallel to margin well beyond the most ventral part of the margin.
Pronotum: relatively short; WH/PW, 0.71; LP/PW, 0.41. Elytra: truncation
not apparent and hence producing a more attenuated appearance than in macu-
losus; epipleural flange absent. Venter: prosternal process of intermediate
length with apex reaching nearly to the posterior line of mesocoxal cavities and
the groove which receives it well posterior to cavities; postcoxal processes
rounded and laterally projecting posterior to midline; outlines of last visible
abdominal sternites of males and females similar with weak nearly symmetrical
crests; not truncated, not produced; sparsely covered with numerous setigerous
punctures. Legs: male pro- and mesolegs scarcely if at all enlarged, and palettes
difficult to distinguish at 20 power magnification; fifth tarsal segment on both
pair of front legs about one and one-half to one and three-fifths as long as cor-
responding fourth; profemoral setae (7 to 8) about equal in size and length to
mesofemoral ones (10 to 12). Genitalia: oval plate asymmetrical with long
produced acuminate tip; its median crest extends anteriorly with little lateral
curvature; raised lateral lines few and weak; aedeagus thin with no distinctive
projections; right paramere elongated and nearly as long as left; each blade or
ovipositor with about 12 rather broadly gapped sawlike teeth.

NOMENCLATURAL NOTES. — | have examined several para-
types of schwarzi, but have not seen the type. No confusion exists
about its identity. The only problem that might be posed is whether
it is a race of undatus or gentilis. Those possibilities seem unlikely;
and in fact, schwarzi seems to share more characteristics with duplex
than any other species. The nearest record of duplex is in Nuevo
Leon, Mexico, which makes it doubtful that the two forms are sub-
specifically related.

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

214 THE GENUS LACCOPHILUS

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This is the rarest species of Laccophilus in the United States and,
perhaps, in North America. It has been collected only in the vicinity
of Washington, D. C., and from Amherst County, Virginia. Fall
(1917) quotes Mr. Sherman as follows: “Originally found by Mr.
Schwarz at Bladensburg, Md., and has been taken so far as I know,
only at the two places (mentioned above), both near Washington,
and only in September, in the almost dried-up brooks, in eddies where
fallen trees or stumps have made dams in which there is an accumu-
lation of dead leaves, etc., and upon the surface of water where there
is a considerable collection of scum and rubbish.” F. N. Young has
also collected them in a stream near Monroe in nearby Amherst
County, Virginia, in late August.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — DISTRICT OF COLUMBIA. District,
3 2 (CARR). MARYLAND. Prince George County. Bladensburg, 13, ix.21.10,
J. D. Sherman (USNM); 2 ¢, 1 2, J. D. Sherman (AMNH). VIRGINIA.
Amherst County. Near Monroe, 2 ¢, 3 @, viii.28.59, F. N. Young (NMSU).
Not located. Ash Grove (near Washington), 15, ix.22.10 (USNM); 9 6,
8 2, ix.22.10; 6 6, 7 2, ix.—.—, J. D. Sherman (AMNH).

Laccophilus duplex Sharp (Figs. 242-249, 328)

Laccophilus duplex Sharp, 1882b, p. 12. Type: male, British Museum (Natural
History), Oaxaca. Zimmermann, 1920, p. 18; Blackwelder, 1944, p. 74.

Laccophilus optatus Sharp, 1882b, p. 13. Type: male, British Museum (Natu-
ral History), Chontales, Nicaragua. Zimmermann, 1920, p. 23; Black-
welder, 1944, p. 74. (New synonym.)

DIAGNOSIS. — This species is very similar to oscillator in general appear-
ance and anatomy. They both have the same general elytral pattern with pale
markings at approximately the same position. The subbasal band appears to be
more restricted in duplex, however, and is more strongly margined by darker
pigment. There is no posterior finger of lighter color from the band in diplex,
as there is in oscillator, nor eroding of the anterior margin of the band in
duplex as in oscillator. L. o. oscillator is larger than duplex, but o. laevipennis
is smaller. Perhaps the best external feature to use is the appearance of the
metacoxal plates. In duplex the plates are nearly yellow except at the lateral
margins where they are darkened to dark brown or nearly black. In contrast
oscillator has dark color suffused throughout the plate, making the venter much
darker. Also, the epipleura of duplex are pale, while in oscillator they are par-
tially darkened. The aedeagus easily separates the two species, however, since
the swollen base of that of duplex is highly diagnostic.

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

216 THE GENUS LACCOPHILUS

L. gentilis suavis and L. ovatus zapotecus are other small forms that have
yellow subbasal markings on the elytra, but they are both smaller and narrower
and have pale metacoxal plates. Both males and females of those two species
have strongly modified last ventral abdominal segments. L. peregrinus may
have yellow longitudinal, instead of horizontal, markings on the elytra, but
either the ovipositor or aedeagus will separate it from duplex.

DESCRIPTION. — Small (length, 3.8 to 4.3 mm; width, 2.1 to 2.3 mm),
brown species with a nearly complete, yellow subbasal elytral band; metacoxal
file absent; prosternal process intermediate; ovipositor sawlike. COLOR. Head:
pale brownish-yellow above and beneath; appendages except mandibles, pale
yellow. Pronotum: generally pale brownish-yellow, but brown along posterior
margin, especially at apex, and some slight darkening near the anterior margin
between the eyes. Elytra: marmorated brown with yellow markings; epipleura
anteriorly translucently pale (may appear to be darker due to darkened margin
of underlying metacoxal plate) and posteriorly reddish-brown. Venter: gener-
ally pale yellowish-brown or brown with a reddish tinge; dark brown or black
around base of mesocoxae and on lateral and hind margins of metacoxal plates.
Genitalia: variable reddish-brown or yellowish-brown. ANATOMY. Méicro-
reticulation: double on head, pronotum and elytra; individual cellules weakly
outlined; surface shining. Head: supraclypeal seam forming two nearly con-
vergent straight lines. Pronotum: relatively short; WH/PW, 0.72; LP/PW,
0.39. Elytra: apices not at all truncated, but tending to be slightly attenuated;
female without epipleural flange (only 20 females examined). Venter: pro-
sternal process reaching an imaginary line drawn across the hind margins of the
mesocoxal cavities and with well-defined ventral crest; postcoxal processes later-
ally scarcely projecting beyond the midline; last abdominal segments not trun-
cated but subtriangular; margins nearly straight in males and slightly convex in
females; median crest asymmetrical in males and slightly convex in females;
both sexes with scattered setigerous punctures and numerous longitudinal rugae
or grooves covering posterior half of the segment; females with marginate
groove extending along margin. Legs: male pro- and mesotarsi only slightly
enlarged and palettes difficult to distinguish at 20 power magnification; fifth
tarsal segment on pro- and mesotarsi only about one and one-half times as long
as corresponding fourth; proleg claws as long as fifth tarsal segment; profemoral
setae (5 or 6) as long as, but finer than mesofemoral ones (6 or 7). Genitalia:
oval plate with produced acuminate tip; its median crest extending anteriorly
without curvature; numerous raised lines on either side; aedeagus with swollen
base and attenuated tip; apex of right paramere short and blunt; ovipositor with
sawlike teeth.

NOMENCLATURAL NOTES. — J. Balfour-Browne has synony-
mized duplex in the collection of the British Museum and the speci-
mens (three cotypes) that I have examined support that conclusion.
Also, F. N. Young has sent me sketches of the elytral patterns and
genitalia of the types of duplex and optatus, and they appear identical.

JAMES R. ZIMMERMAN ANG

Sharp’s descriptions differ significantly only in the statement that du-
plex has a prosternal process not elongate and that optatus has the
prosternal process elongate. L. duplex was described from Oaxaca
and optatus from Nicaragua, and this appears to be the principal basis
for separating them. He remarks that duplex resembles the Brazilian
L. fumatus Sharp and optatus is closely allied to L. latifrons Sharp.
I have not seen either of these species.

VARIATION. — The yellow subbasal markings vary from a com-
plete band with little interruption, even at the elytral suture, to three
separate discrete marks. The difference is geographical, since the
Honduras specimens tend toward discreteness and the Mexican ones
toward continuity. There is no obvious difference in size between the
sexes (fig. 24).

The individuals from San Luis Potosi are significantly larger than
those from Costa Rica (Table 23). The three males from Chiapas
are closer to the size of the southern population. There is enough
difference in pattern and size between the populations from northeast-
ern Mexico and those from below the Isthmus of Tehuantepec to
warrant description of two races, but it will have to wait until more
is known about the limits of each.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This species ranges from near Monterrey in Nuevo Leon to Brazil
(fig. 23). It occurs at low altitudes, but above the coastal plain.
I have collected it in heavily shaded, shallow, clear streams with
gravel bottoms and in roadside ditches.

MATERIAL EXAMINED

COSTA RICA. — Esparta, 8 m. NW., 1 2, vi.26.67; Puntarena, 1 6, 4 9,
vii.22.65; Liberia, 8 m. SE., 21 6, 31 2, vii.24.65, P. J. Spangler; San Jose,
1 4,1 2, vii.16.57, D. Lauck (USNM).

EL SALVADOR. — Cd. Arce, 7 m. SE., 2 2, viii.3.65, P. J. Spangler (USNM).

HONDURAS. — Ruatan I., 1 6 (BM). San Marcus Colon, 1 6,4 92, vii.28.65;
N. Sabina Grande, 4 4, 2 2, vii.29.65, P. J. Spangler (USNM).

MEXICO. — CHIAPAS. Chiapa de Corzo, 1 6, ix.1.63; Cuahtemoc, 3 6,
viii.30.63, JRZ (NMSU). NUEVO LEON. Linares, 20 m. W., Linares R.,
1 6, xi.16.48, H. B. Leech (CAS). Monterrey, 4 m. S., 1 46, vii.7.63, Arnett
& Van Tassel (CUA). Santiago, 4m. N., 1 é, vii.8.63, JRZ (NMSU). SAN
LUIS POTOSI. Cuidad del Maiz, 15 m. E., 1 6, xi.19.48, H. B. Leech (CAS).
El Salto, 1 @, iii.25.63, JRZ (NMSU). Palitla (5S m. N. of Tamazunchale),

MEM. AMER. ENT. SOC., 26

218 THE GENUS LACCOPHILUS

2 6,5 92, xii.22.48, H. B. Leech (CAS). Tamazunchale, 1, vi.30.48, Werner,
Nutting (ARI); Quinta Chilla, 3 ¢, 9 9, xii.21.48 (CAS). TAMAULIPAS.
Antiguo Morelos, 3 m. N., 1 2, iii.26.63; Ocampo, 2 @, iii.24.63, JIRZ (NMSU).
VERACRUZ. Near Tinaja, 1 @, viii.25.62; Papantla, 18 km. E., 1 2, ix.9.64,
JRZ (NMSU).

NICARAGUA. — Chontales, 2 ¢ (BM).

LACCOPHILUS OSCILLATOR

Sharp (1882b) described oscillator and laevipennis as separate
species, recognizing that they were closely related. In fact, he said
the markings of the two were identical, but that they differed in size
and in microsculpture. I find no difference in the last feature, but
they differ in size. One was described from Oaxaca and the other
from Guatemala. It is now known that the range of these two extends
through Central America and up the Sierra Madre Occidental into
Arizona. Intergrades occur in Nayarit, Jalisco, and Michoacan.
Oddly enough, the range of the northern race, oscillator, extends all
the way to Oaxaca, but apparently at higher elevations than does
laevipennis.

The aedeagi of the two differ only in size, and all other sexual
characters that might be diagnostic are similar. In fact, except for the
unusual elytral patterns and the darkened venter, it is difficult to point
out a useful diagnostic character for the species.

DESCRIPTION. — Small to medium (length, 3.5 to 5.8 mm; width, 2.4 to
3.4 mm), marmorated brown species with a nearly complete subbasal elytral
fascia; metacoxal file absent; prosternal process short; ovipositor sawlike.
COLOR. Head: pale brownish-yellow above and beneath except for a variable
amount of reddish-brown on the occiput between the eyes; appendages except
mandibles pale. Pronotum: generally pale brownish-yellow, but with a variable
amount of reddish-brown on the disc, anterior margin, and apex; posterior
margin translucently reddish-brown. Elytra: marmorated smoky brown with
yellow markings, forming a nearly complete subbasal band and a much less
complete postmedian one; the latter composed of three or four elongate marks
which are generally, but unevenly arranged in an anterior-posterior direction;
epipleura partially darkened anteriorly as well as posteriorly. Tergite VIII:
proximally brown, but distally pale yellowish-brown. Venter: prosternum, pro-
and mesolegs pale yellowish-brown with reddish tinge; remainder variable from
yellowish-brown to sometimes suffused with reddish-brown or black; nearly
black around the bases of the mesocoxae, the outer half of the metacoxal plates,
and on parts of abdominal segments; first visible abdominal segment usually
pale in contrast to those posterior. Genitalia: variably pale yellowish-brown to

JAMES R. ZIMMERMAN 219

~ \} LACCOPHILUS 0. OSCILLATOR o

INTERGRADE 0°

LACCOPHILUS O. LAEVIPENNIS @

Figure 26. Distribution of Laccophilus oscillator.

dark brown. ANATOMY. Méicroreticulation: double on head, pronotum, and
elytra; cellules of inner meshes very weak. Head: supraclypeal seam nearly
parallel to margin, but may arch slightly upward near midline. Pronotum:
WH/PW, 0.70; LP/PW, 0.39. Elytra: apices not truncated; female epipleura
without flange. Venter: prosternal process short; postcoxal process neither
rounded nor produced laterally, but forming a nearly straight line across the
posterior margins; last visible segment not truncated in either sex and similar;
male segment slightly produced; its median crest frequently indistinct; posterior
marginated grooves in males and females not attaining the apex; numerous
rugae and prominent setigerous punctures also present. Legs: male pro- and
mesotarsi scarcely enlarged; fifth tarsal segment on both pair of front legs about
twice as long as corresponding fourth; palettes difficult to distinguish at 20
power magnification; profemoral marginal setae (5 to 7) as long as, but finer
than mesofemoral ones (8 to 11). Genitalia: oval plate relatively small and
notched on either side of the acuminate produced tip; its ventral median crest
pronounced and extending anteriorly with little or no lateral curvature; numer-
ous fine raised lines on either side of the crest; aedeagus curving to a small
tapered apex; right paramere nearly as long as left; ovipositor with about 10 or
11 sawlike teeth.

MEM. AMER. ENT. SOC., 26

220 THE GENUS LACCOPHILUS

o. oscillator Uruapan, Michoacan

3.4 3.6 3.8 4.0 4.2 4.4 4.6

o. oscillator Arizona

3.4 3.6 3.8 4.0 4.2 4.4 4.6

intergrades Autlan, Jalisco

3.4 3.6 3.8 4.0 4.2 4.4 4.6

La Resolana, Jalisco

l rare , 72

3.4 3.6 3.8 40 4.2 4.4 4.6

NUMBERS

laevipennis

La Huerta, Jalisco

3.4 3.6 3.8 4.0 4.2 44 4.6
TOTAL LENGTH (mm)

Figure 27. Histograms of length in Laccophilus oscillator. Males shown
in crosshatch; females, stippled.

221

ZIMMERMAN

JAMES R.

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MEM. AMER. ENT. SOC., 26

D22, THE GENUS LACCOPHILUS

NOMENCLATURAL NOTES. — The only apparent problem
with this race is in its relationship to laevipennis. Since intergrades be-
tween the two have been found, the problem has resolved itself. Sharp’s
cotypes of oscillator and laevipennis may be confused with each other,
but the types themselves are easily distinguishable and seem to repre-
sent clearly each of the separate races.

Laccophilus oscillator oscillator Sharp, new status (Figs. 250-257, 326)

Laccophilus oscillator Sharp, 1882b, p. 11. Type: British Museum (Natural
History), male; Oaxaca (Hoege). Zimmermann, 1920, p. 23; Blackwelder,
1944, p. 74.

DIAGNOSIS. — Elytral pattern, dark venter, and lack of a metacoxal file
serve to separate this race from all other forms but /aevipennis, which is smaller.
The non-irrorated elytral pattern is quite similar to those found in duplex and
subsignatus Sharp; but both of those have only a suggestion of dark color on the
margins of the metacoxal plates, while o. oscillator has its plates strongly dark-
ened over most of their surface. The pattern of duplex has a more frothy ap-
pearance as well. L. subsignatus appears to occur only as far north as Panama
which is well below the range of o. oscillator. The latter form is larger as well
(Table 24). L. horni, L. peregrinus, L. gentilis, L. leechi, L. pictus, and L.
youngi are sympatric with o. oscillator and have non-irrorated patterns; but
each differs considerably in detail and color.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
—L. o. oscillator has a long, narrow range. It extends from southern
Arizona in Pima and Santa Cruz counties to the valley of Oaxaca in
the Sierra Madre Occidental, Central Volcanic Highlands, and the
Sierra Madre Sur (fig. 26). It appears to be taken most frequently
from about 4000 to 6000 feet. From Nayarit south L. o. oscillator
appears to intergrade with o. laevipennis below 4000 feet. It inhabits
pools of rocky mountain streams that are frequently subjected to tor-
rential currents. In fact, L. oscillator seems to be one of the very few
Laccophilus that prefers current to pools. I have collected them by
sweeping on the under side of large rounded boulders that are subject
to heavy current. The race often occurs with L. horni and L. pictus.

Collections have been spotty, but this is undoubtedly due to the
relative inaccessibility of its usual habitat and not due to any real gap
in its range.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — ARIZONA. 4 2,2 ¢ (ANSP). Pima
County. Sta. Catal. Mtns., Molina Basin, 1 9, vi.11.58, C. & G. MacNeill

JAMES R. ZIMMERMAN 223

(CAS); 1, iv.18.53; 1 2, A. & H. Dietrich (CNL); Bear Canyon, 1 9, vi.18.53,
A. & H. Dietrich (CNL). Santa Cruz County. Pena Blanca, Pajarito Mtns.,
1 9, vi.28.62; 1 2, vii.20.62, Arnett & Van Tassel (CUA). Santa Rita Mtns.,
Florida Can., 1 2, iv.19.59, F. G. Werner (ARI); 1 ¢@, ix.7.61; 4 6, 17 9,
xii.2.61, JRZ (NMSU).

MEXICO. — JALISCO. Colima, 29 m. NE., 1 6, 3 2, xii.3.48; Mazamitla,
1 9, xii.1.48; 17 m. S., 2 2, xii.5.48, H. B. Leech (CAS). MICHOACAN.
Tzitzio, 1 2, vii.27.62; Uruapan, 10 m. S., 3 6, 2 2, ili.26.64, JRZ (NMSU).
OAXACA. Huajuapan, 17 km. NE., 1 2, ix.3.64, JIRZ (NMSU). Tutla, 1 ¢,
1 9, xii.13.48, H. B. Leech (CAS). Oaxaca, 3 m. S., 1 4, viii.26.63, JRZ
(NMSU). SONORA. MHuachinera, 3 2, vii.25.63, JRZ (NMSU).

Laccophilus oscillator laevipennis Sharp, new status
(Figs. 258, 259, 327)

Laccophilus laevipennis Sharp, 1882b, p. 12. Type: British Museum (Natural
History), male; San Joaquin, Guatemala City (Champion). Zimmermann,
1920, p. 23; Blackwelder, 1944, p. 74.

DIAGNOSIS. — Except for its smaller size, the same diagnostic characters
used for o. oscillator apply to o. laevipennis. Within its range it will most
probably be confused with duplex, leechi, g. suavis, ovatus, and subsignatus.
It differs from duplex in having a dark venter, from /eechi in the elytral pat-
tern and a lighter pronotum, from suavis by lack of a metacoxal file and dark
venter, and from ovatus by elytral pattern, dark venter, and simple margin of
the last ventral segments in either sex. L. subsignatus appears to extend no
farther north than Panama, and laevipennis may go no farther south than Costa
Rica. They are extremely similar, however. At this time, it is not known if
they are races of the same species.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— L. o. laevipennis has been collected from Jalisco to Costa Rica (fig.
26). Collections are also sparsely distributed in Oaxaca, Chiapas,
and Nicaragua. It occurs in mountainous habitats in tropical decidu-
ous forest. It is found in the same kind of habitats as o. oscillator.

MATERIAL EXAMINED
COSTA RICA. — San Jose, 1 6, vii.16.57, D. Lauck (USNM).

MEXICO. — CHIAPAS. Simojovel, 3 ¢, 1 9, viii.6.58, J. A. Chemsak (BERK).
Cuahtemoc, | 4, viii.30.63; Chiapa de Corzo, 3 ¢, 3 9, viii.28.63, JRZ (NMSU).
JALISCO. Colima, 29 m. NE., 2 6, 1 2, xii.3.48, H. B. Leech (CAS). La
Huerta, 6 m. S., 8 6, 4 2, xi.26.66; 9 m. S., 41 6, 40 2, x.26.66, A. Smith &
JRZ (NMSU). OAXACA. Tapanatepec, 1 2, xii.31.55, J. C. Schaffner
(UMMZ). Near El Cameron, Rd. 190, 2 6, viii.27.63, JRZ (NMSU).

MEM. AMER. ENT. SOC., 26

224 THE GENUS LACCOPHILUS

NICARAGUA. — San Benito, 13 m. N., 3 2, vii.11.65; Somoto, 1 2, vii.28.65;
Esteli, 9m. N., 1 @, vii.10.65, P. J. Spangler (USNM).

INTERGRADATION IN LACCOPHILUS OSCILLATOR

The only appreciable difference between the two races of L. oscil-
lator is size; and there is, apparently, almost no overlap in that char-
acter. The northern race is nearly a millimeter longer (Table 24).
Elytral pattern, color, and sexual characters are much the same.
Populations in Nayarit, Jalisco, and Michoacan are clearly intermedi-
ate, however.

It is possible to show an almost complete conversion in less than
fifty miles from a pure population of laevipennis to a nearly pure one
of oscillator (fig. 27). The largest individuals barely exceed 3.9 mm,
while unmixed populations of oscillator from Arizona and Michoacan
have none or less than 4.1 mm. Although the data is conflicting,
males and females appear to be about the same size in both races; and
thus, the discussion can be simplified. The La Huerta sample has
steep slopes and a well-defined peak when one plots the frequency
distribution of length. Standard deviations are also relatively low
(Table 24). Both are indicative of a homogenous population. Two
other samples, from inland of that locality, show an interesting progres-
sion to a population that is predominately oscillator. Other samples
in Nayarit and Michoacan represent a continuation toward oscillator.
About twenty miles northeast of La Huerta near La Resolana (Casi-
miro Castillo), the population has a bimodal distribution with one
peak at the same length as for /aevipennis at La Huerta; but there is a
second less definite one at about 4.2 mm which is closer to the mean
for oscillator. Also, a few individuals fall in the intermediate size
range of 3.9 to 4.1 mm. Continuing on to nine miles southwest of
Autlan, the average length of the population has shifted to about 4.3
mm; and only a small percentage of the population is in the laevi-
pennis size range. At this locality the size separation of the two races
is almost completely obscured. Unfortunately, I do not have a sample
from just a few more miles inland that would demonstrate a complete
shift to oscillator. One must interpolate between the Autlan sample
and those in Arizona and Michoacan. There are two samples from
Michoacan. One, at Tzitzio, shows intergradation; but the other, at
Uruapan, does not. The samples from Nayarit and Tzitzio are very
similar to the one from Autlan.

JAMES R. ZIMMERMAN DHS)

The zone between the two races is probably due to secondary
contact. Although there is a uniform gradation in size between them,
the zone itself is extremely narrow. Ecological differences appear to
be slight. Obviously, oscillator can tolerate cooler temperatures; but
they are both found in rapidly flowing, shaded mountain streams.

MATERIAL EXAMINED
L. o. oscillator X L. o. laevipennis

MEXICO. — JALISCO. Autlan, 9 m. SW., 36 6, 27 2, x.24.66; La Resolana
(Casimiro Castillo), 3 m. N., 71 6, 57 2, x.26.66, A. Smith & JRZ (NMSU).
MICHOACAN. Tzitzio, 3 m. S., 7 6, 4 9, vili.22.63, JRZ (NMSU). NA-
YARIT. Tepic, 25 km. S., 1 9, ix.24.53, B. Malkin; Sierra de Zapotan, | ¢,
6 2, xii—.42, E. Paredes (CAS).

LACCOPHILUS GENTILIS

This polytypic species is composed of three allopatric populations
which are considered as two subspecies. L. gentilis gentilis LeConte
is known from Florida to Louisiana. L. g. suavis Sharp occurs from
Tamaulipas to Campeche and Costa Rica in a continuous series of
populations. Another population extends from Colima to Sonora on
the western side of the continent. It is unlikely that there is any real
geographical separation between the last two groups, since there has
been no intensive collecting for Laccophilus in the coastal regions of
Michoacan, Guerrero, or western Oaxaca.

This species presents an interesting situation in the North Ameri-
can Laccophilus species, since the nominate race has virtually lost the
metacoxal file that is still present in swavis. The species is well repre-
sented in the West Indies, but those populations are not considered
here.

DESCRIPTION. — Small (length, 3.2 to 3.7 mm; width, 1.7 to 1.9 mm),
reddish-brown species with or without a yellow, subbasal elytral fascia; meta-
coxal file present or absent; prosternal process long; ovipositor sawlike COLOR.
Head: reddish-brown or yellowish-brown; appendages of matching color. Pro-
notum: generally yellowish-brown laterally, but much darker anteriorly between
the eyes and posteriorly across the apical angle; discal region may be dark-
ened to a lesser degree. Elytra: generally reddish-brown or brown with paler
brownish-yellow or reddish-brown areas, especially subbasally; anterior part of
epipleural pale, posterior darkened. Tergite VIII: pale yellowish- or reddish-
brown. Abdomen usually darker. Genitalia: same as venter. ANATOMY.
Microreticulation: double on head, pronotum, and elytra; primary mesh weak;
surface shining. Pronotum: anterior margin covering the back of the eyes;

MEM. AMER. ENT. SOC., 26

226 THE GENUS LACCOPHILUS

sides less convergent anteriorly than in most Laccophilus,; WH/PW, 0.73;
LP/PW, 0.38. Elytra: apices slightly truncated; female epipleura rarely, if
ever, with flange. Venter: prosternal process long, extending beyond margin
of mesocoxal cavities; metacoxal file present and of about 8 or 9 ridges in males
and females of suavis, but these only suggested in g. gentilis; postcoxal processes
nearly straight across, with lateral parts barely extending past the midline; last
visible segment of males and females truncated; females less so and with emargi-
nate apex; weak crest on male segment; a small tubercle on either side; females
with fairly acute crest; in both sexes a row of thick, short hairs or setae on the
lateral margins of segment. Legs: pro- and mesotarsi scarcely enlarged in a
dorsoventral plane; fifth tarsal segment about twice as long as corresponding
fourth on both pair of front legs; palettes difficult to distinguish at 20 power
magnification; profemoral marginal setae (4 or 5) shorter and finer than meso-
femoral ones (5 or 6). Genitalia: oval plate with produced acuminate tip;
strongly asymmetrical; its ventral crest well developed, but interrupted about
half the length of the plate; no raised lines on either side of crest; aedeagus with
a crest on its convex side and resembling a bird’s head in profile view; para-
meres relatively short, especially the left one when compared to the aedeagus;
ovipositor with about 10 pair of sawlike teeth.

Laccophilus gentilis gentilis LeConte, new status (Figs. 260-267, 331)

Laccophilus gentilis LeConte, 1863, p. 23. Type: Museum of Comparative
Zoology, number 5975, female, Wapler (collector?), Louisiana; Crotch,
1873, p. 400; Sharp, 1882a, p. 300; Leng and Mutchler, 1918, p. 78;
Blatchley, 1919, p. 308; Leng, 1920, p. 77; Zimmermann, 1920, p. 19;
Blatchley, 1932, p. 302; Young, 1954, p. 47.

DIAGNOSIS. — The largest individuals do not attain 4.0 mm, which makes
this the smallest Laccophilus in the United States. L. g. gentilis, L. undatus,
and L. schwarzi are all non-irrorated, but the last two exceed 4.0 mm and occur
no farther south than Virginia. Both of these species have prominent yellow
elytral markings that are lacking on this race of gentilis. The elongate proster-
nal process and spines on last ventral segment will also separate g. gentilis from
any other United States species. The principal difference between g. gentilis
and g. suavis is that a file and yellow subbasal elytral band are present in the
latter and not in the former.

NOMENCLATURAL NOTES. — There has been no confusion
of gentilis with other species. The close relationship between it and
suavis has not been previously recognized. The two differ in elytral
patterns and in the presence of a file in one, but not the other. In
both traits there is evidence of relationship, however. The same basic
pattern is present in both, but the lighter areas are reddish in gentilis
and usually yellow in suavis. There is a trace of the file in gentilis,

227

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

228 THE GENUS LACCOPHILUS

if one examines the metacoxal plate carefully; and the file is weak in
suavis and has been reduced to less than ten lines. Sexual characters
are much the same in both. There is no test of sympatry, since their
ranges are widely separated. More intensive collecting in coastal
regions of Tamaulipas and Texas might show connecting and inter-
grading populations, however. It seems fairly certain that this species
has representatives on several Caribbean islands, but I have not exam-
ined enough material to make nomenclatural conclusions about those.

VARIATION. — There is considerable individual variation in the
degree to which the irregular reddish markings occur as spots or bands
extending over the surface of the elytra. The reddish markings corre-
spond to yellow ones in suavis. There is also some variation in the
amount of darkening present upon the pronotum.

Females are slightly larger than males (Table 25). From a sam-
ple of 22 females and 11 males, the means for total length were 3.46
and 3.38. The largest female in the sample is 3.64 mm long, and the
largest male is 3.48 mm long.

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— Young (1954) states that gentilis occurs in Florida almost through-
out the peninsula, but not west of the Apalachicola River (fig. 23).
There are also two records from southern Georgia and one from Lou-
isiana. The type was described from Louisiana. Its known range is
well removed from the nearest record of suavis, which is southern
Tamaulipas.

Young says that in northcentral Florida it is most abundant in
sloughs or swamps along the edges of lakes. It more rarely is found
in small streams and various kinds of ponds. I have not collected this
subspecies.

MATERIAL EXAMINED

UNITED STATES OF AMERICA. — FLORIDA. Alachua County. New
Man’s Lake, 5, iv.12.25, M. D. Leonard (CML). Gainesville, 2 3, 2 9,
vi.23.59, R. E. Woodruff (MCZ). Dade County. 5, vii.18.38 (CNL). Home-
stead, 1 2, vii.27.57, R. M. Baronowski (CAS); 1 ¢, 2 9, vi.—.29, Darlington
(MCZ). Duval County. Jacksonville, 1, 8—.02 (USNM). Flagler County.
Crescent City, 2 (USNM). Hendry County. LaBelle, 2, ii.26.18, W. S. Blatch-
ley (USNM). Hernando County. Croom, 6 @, vi.18.29, P. J. Darlington
(MCZ). Highlands County. Brighton, 5 ¢, 4 2, vi.16.29, Darlington (MCZ).
Boundary of Okeechobee County & Glades County, Mouth of Kissimmee River,
1, iv.6.39, J. C. Bradley (CNL). Orange County. Winter Park, 1 6, vii.8.40,

JAMES R. ZIMMERMAN 229

H. T. Fernald (MCZ)_ Pinellas County. Dunedin, 2, iv.17.25; 1, 1.16.18;
LISS eexcle li weleiie2 S84 2. ively. We S Blatchleys (USNM):
2, 1.22.22 (CNL). Sarasota County. Sarasota, 1 6, 2 9, ——.11, W. S.
Blatchley (MCZ). Taylor County, 1 (USNM). GEORGIA. Charlton County.
Okefenokee Swamp, Bog, 1, xii.26.13 (CNL). Jefferson County. Wrens, 1,
ii.8.11 (CNL); 1 (ANSP). LOUISIANA. Vermillion County. Gueydan
(light), 1, vii—.25, E. Kalmbach (USNM). Gueydan, 1, vii.11.25, Wapler
(collector?), 1 2 (MCZ).

Laccophilus gentilis suavis Sharp, new status (Fig. 332)

Laccophilus suavis Sharp, 1882b, p. 13. Type: British Museum (Natural His-
tory), male, Cordova, Veracruz; Zimmermann, 1920, p. 26; Blackwelder,
1944, p. 74.

Laccophilus championi Sharp, 1882b, p. 14. Type: British Museum (Natural
History), male, Paso Antonio, Guatemala; Zimmermann, 1920, p. 17;
Blackwelder, 1944, p. 74. (New synonym.)

DIAGNOSIS. — This is the only North American Laccophilus that has a
metacoxal file and a long prosternal process. L. ovatus has an equally long
process, but no file. The two are also about the same size (average length
about 3.5 mm) and have similar reddish-brown elytra with yellow subbasal
markings that form a complete band in suavis, but may be interrupted in ovatus.
The reddish-brown pigment is more uniformly applied in swavis than in ovatus.
L. duplex and L. oscillator have similar patterns, but are larger species (about
4.0 mm or longer), lack files, have shorter prosternal processes, and have un-
truncated last ventral abdominal segments. The last ventral segments of suavis
and ovatus are similarly ornamented with lateral spines, small protuberances,
and emarginate apices in the females. L. g. suavis males and females lack the
prominent produced apices of ovatus. The aedeagi of the two species are quite
different, however.

NOMENCLATURAL NOTES. — The reason for reducing suavis
was discussed under gentilis, but it is also necessary to synonymize
championi Sharp with suavis. The former was described from Guate-
mala and the latter from Cordova, Veracruz. I have examined cotypes
(= paratypes) of both from those localities and find the differences to
be individual and not geographical. Page precedence establishes the
priority of suavis.

VARIATION. — The extent of the yellow on the subbasal band,
the midlateral spot, and the postmedian markings are the most appar-
ent variations in color and pattern. There is always some yellow or
reddish-yellow color at all of these positions, but considerable reduc-
tion sometimes occurs. Some specimens have darker, more extensive
pronotal marks than others, with darkening reaching clear across the

MEM. AMER. ENT. SOC., 26

230 THE GENUS LACCOPHILUS

disc. The reddish-brown on the elytra may darken in some to very
dark brown. Specimens from the west coast of Mexico are darker
reddish-brown than those from Veracruz, with more contrast between
light and dark areas.

As in gentilis, the females appear to be larger than the males.
There is close agreement also in the populations from Veracruz and
those from Nayarit (Table 25). The EL/PW ratio varies from 0.538
to 0.550. This race is larger than gentilis (fig. 24).

GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— This is a tropical lowland Laccophilus. Its range is probably con-
tinuous from southern Tamaulipas on the northeast and southern So-
nora on the northwest to El Salvador or farther into Central America
(fig. 23). The large gap between Colima and the Isthmus of Tehu-
antepec is undoubtedly due to a lack of collecting. On the east coast,
it has a gap between southern Tamaulipas and central Veracruz, but
this is also another poorly collected area.

It was mostly collected in still water on the coastal plain. It occurs
in grassy-bottomed or mud-bottomed pools that form in roadside exca-
vations, or in duckweed-choked, shaded sloughs, or in pools of stabi-
lized dunes immediately next to the ocean. On a few occasions, it
was at several thousand feet, however. There is one record near
Zamora, Michoacan (over 5000 feet), and several from the vicinity
of Tepic, Nayarit (3000 feet), which indicate it readily migrates over
wide areas. I have not yet been able to collect suavis in large num-
bers, probably because of not finding its preferred habitat.

MATERIAL EXAMINED

MEXICO. — CAMPECHE. Champoton, 19 m. S., 10 6, 14 2, vii.26.64,
P. J. Spangler (USNM); 11 m. W., 1 6, 1 2, xi.27.63; Cuidad de Carmen,
4m.E., 1 ¢, xi.27.63, JRZ (NMSU). CHIAPAS. Cintalapa, 5 m. W., 1 ¢,
ix.1.63, JRZ (NMSU). COLIMA. Manzanillo, 5 m. S., 5 ¢, 1 2, vii.29.62,
JRZ (NMSU). JALISCO. Barra de Navidad, 2 ¢, 1 9, 1.22.61, C. O. Morse
(CAS); 1 6,5 2, x.25.66; La Huerta, 1 6, x.25.66, A. Smith & JRZ (NMSU).
NAYARIT. Pena, 12 m.N., 1 2, xi.26.48, H. B. Leech (CAS). Tepic, 1 6,
1 2, ix.15-17.53; 1 6, 1 Q, ix.21-24.53, B. Malkin; 5 m. SE., 2 9, ix.26.48,
H. B. Leech; San Blas, 1 ¢, ix.28.61, C O. Morse; 2 ¢, 1 9, xi.7.58, I. Moore
(CAS);5m.E., 7 &, 10 @, vii.31.62, JIRZ (NMSU). MICHOACAN. Zamora,
9m. W., 1 6, xii.6.48, H. B. Leech (CAS). OAXACA. Tehuantepec, 1 6,
1 2, ix.6.64, JRZ (NMSU). QUINTANA ROO. Cozumel Island, Celarain
Point, 1 4, iv.21.60; San Miguel, 2 m. N., 1 6, 3 2, iv.3.60, J. F. G. Clarke

JAMES R. ZIMMERMAN 231

(USNM). SINALOA. Mazatlan, 1 4, viii.14.60, Arnaud, Ross, Rentz (CAS);
7m. S., 1 ¢, xii.11.62, JRZ (NMSU). Pericos, 26 m. N., 1 ¢, viii.13.60,
Arnaud, Ross, Rentz (CAS). TABASCO. Villahermosa, 1 46, 3 92, vii.25.64;
23 m. N., 1 6, 4 Q, vili.6.64, P. J. Spangler (USNM); 5 m. S., 2 9, x1.26.63,
JRZ (NMSU). TAMAULIPAS. Mante, 1 é, iii.23.63,JRZ (NMSU). VERA-
CRUZ. Acayucan, 20 m. S., 1 6, ix.7.64; Alvarado, 10 m. W., 5 6, 6 2,
JRZ (NMSU). Cuitlahuac, 1 ¢, 2 92, viii.10-12.64, P. J. Spangler (USNM).
Jalapa, 10 m. E., 9 6, 4 9, vii.27.62, JRZ (NMSU). J. D. Covarrubia, 1 m.
N., 5 6, 6 Q, viii.26.62, P. J. Spangler (USNM). Near Garro, 2 6, 6 2,
ix.8.64; Santiago Tuxtla, 10 km. S., 1 6, 1 92, ix.8.64, JRZ (NMSU). Tres
Zapotes, 1 6, iv.11.39, A. Wetmore (USNM). Veracruz, 14 m. SE., 1 4,
vi.16.58, J. C. Schaffner (UMMZ). Cordova, 1 6, cotype, B. C. A. Coll.
(BM).

BRITISH HONDURAS. — 1 ¢, —.—.05 (BM).

EL SALVADOR. — Acajutla, 19 m. W., 1 4, vii.8.65; La Union, 15 m. SW.,
26,4 9, vii.31.65; Metalio, 1 3, 4 9, viii.4.65; Cd. Arce, 7 m. SE., 4 6, 6 9,
vili.3.65, P. J. Spangler (USNM).

CUBA. — Camaguey, Baragua, | 6, vi.5.32, Christenson; Cayanas, | 4, viii.5.08;
1 6, ix.5.—; 1 2, 1.6.—, E. H. Schwarz (USNM).

COSTA RICA. —1 2, 1.8.28; 1 9, iii.26.36, F. Nevermann; Taboga Agr.
Exp. Sta., 2 ¢, 3 2, vi.27.67, P. J. Spangler (USNM).

GUATEMALA. — Paso Antonio, 2 ¢, B. C. A. Coll. (BM).
PANAMA. — Gamboa, 4 m. W., 1 9, vii.6.67, P. J. Spangler (USNM).

Laccophilus ovatus zapotecus, new subspecies (Figs. 268-275, 330, 331)

DIAGNOSIS. — This race differs from the one in South America in the
aedeagus and in the presence of a nearly complete subbasal elytral band instead
of a few, indistinct yellow markings at that position. The aedeagus of the
nominate race is more gracefully ballet-footed; L. 0. zapotecus differs from
L. gentilis suavis in the absence of a file in the male and in the slightly differ-
ent shape of the male and female last ventral segments. L. ovatus is more
parallel-sided, due to the longer widened anterior portion of the female epi-
pleura. L. duplex, L. oscillator, L. gentilis, suavis, and L. ovatus zapotecus
have subbasal bands, mid-lateral spots, and several post-median marks on the
elytra; but oscillator, and sometimes duplex, are dark beneath and both longer
and broader. The patterns of zapotecus and g. suavis are very similar, but that
of zapotecus is more unevenly marmorated, and has a greater concentration of
dark pigment next to the yellow marks. Sexual characters are better than pat-
tern for separating these two sympatric species.

DESCRIPTION. — Small (length, 3.25 to 3.75 mm; width, 1.75 to 1.95
mm), reddish-brown subspecies (and species), marked with yellow subbasal
elytral band; metacoxal file absent, prosternal process long; ovipositor sawlike.

MEM. AMER. ENT. SOC., 26

232 THE GENUS LACCOPHILUS

COLOR. Head: entirely pale brownish-yellow (sometimes tinged with red)
above and beneath. Pronotum: generally same as the head, but anteriorly a
pair of merged reddish-brown markings and a darker mark at the posterior
apex; entire posterior margin translucently reddish-brown. Elytra: marmorated
reddish-brown with pale yellowish-brown markings; anterior half of the epi-
pleura pale, posterior half darkened. Tergite VIII: brown. Venter: entirely
pale brownish-yellow, sometimes tinged with red, especially the hind legs. Geni-
talia: variably reddish-yellow brown. ANATOMY. Méicroreticulation: double
on head, pronotum, and elytra with individual cellules weakly impressed; sur-
face shining. Head: supraclypeal seam arching slightly upward at midline.
Pronotum: WP/PW, 0.73; LP/PW, 0.39. Elytra: apices slightly truncated;
female epipleura not narrowed until level of posterior margin of second ab-
dominal segment instead of anterior to the posterior margin of the first segment.
Venter: prosternal process long with apex extending beyond imaginary line
drawn across posterior margins of hind coxae; process with well-defined crest;
postcoxal processes truncated along the posterior margins, but emarginate at
the midline; last abdominal segment in both sexes truncated but with produced
apex; female apex emarginate with ventral crest; males with asymmetrical irregu-
lar median crest and a darkened, sclerotized oblique ridge on the left side; both
sexes with multiple rows of thick, short spines on the lateral margins of the last
three abdominal segments of the venter and on the female valves (Sternite
VIII) on either side of the ovipositor. Legs: pro- and mesolegs enlarged in a
dorsoventral plane; fifth segment about one and two-thirds as long as corre-
sponding fourth; claws nearly as long as its corresponding fifth segment; palettes
easily visible at 20 power magnification; profemoral setae (6) nearly as long,
but thinner than mesofemoral ones (6 or 7). Genitalia: oval plate with pro-
duced acuminate tip; its ventral crest raised, well-defined, curving slightly to the
left anteriorly; some weak raised lines on the right side; aedeagus flattened with
profile resembling a dancer’s leg; right paramere with produced apex and nearly
as long as left; ovipositor with about 14 very fine sawlike teeth.

NOMENCLATURAL NOTES. — While this race is similar to
the Brazilian population, there are several differences which support
subspecific rank. The range is too incompletely known, however, to
determine whether they are allopatric populations, or if there is an
intergrade zone between them, or if they overlap as separate species.

VARIATION. — The most obvious variation in pattern is the
extent to which the elytral subbasal band is interrupted. In Tabasco
specimens the band is complete; but in some in Chiapas, there are
three discrete marks which is an intermediate condition to the only
specimen (a male cotype) from Brazil that | have examined.

Males and females seem to be about the same length (Table 26,
fig. 24). A total of 23 females had a mean length of 3.53 mm, and

233

JAMES R. ZIMMERMAN

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MEM. AMER. ENT. SOC., 26

234 THE GENUS LACCOPHILUS

one of 19 males had a mean of 3.58 mm. There is little difference in
the values from three localities; one in Veracruz, another in Tabasco,
and the third in Chiapas. Females may be slightly broader than males,
however; and this is indicated by PW/EL ratio. The one Brazilian
specimen that I have examined had a length of only 3.25 mm, which
is smaller than the smallest individual from the Mexican localities.
GEOGRAPHICAL RANGE AND HABITAT PREFERENCES.
— The northern limit of the range of this race is about the Isthmus of
Tehuantepec. The southern limit is not known, but it reaches Panama
(fig. 17). All localities are under 500 feet elevation. It was found
mostly in pasture ponds and in roadside ditches with mud bottoms.

MATERIAL EXAMINED

Holotype male, allotype female, and a male and female paratype with the
following locality data are in the United States Nation! Museum: 5 miles south
of Villahermosa, Tabasco, xi.26.63, J. R. Zimmerman. A male paratype with
the same data as the holotype is in the California Academy of Sciences, San
Francisco, and a male and female with the same data are each in the University
of Michigan Museum of Zoology, Ann Arbor, and the Departmento de Ento-
mologia, Laboratorio de Sanidad Vegetal, Coyoacan, D. F., Mexico. Another
female paratype from one mile north, J. D. Covarrubia, Veracruz, viii.26.62,
J. R. Zimmerman, is also in the California Academy of Sciences.

COSTA RICA.— Toboga Agr. Exp. Sta., 4 6, vi.27.67; 1 6, 2 2, v.28.67,
P. J. Spangler (USNM). La Cruz, 16 m.S., 3 6, 1 2, vii.13.65, P. J. Spangler
(USNM).

GUATEMALA. — Paso Antonio, 2 2, Champion, B. C. A. Coll. (BM).
NICARAGUA. — Rivas, 22 m. S., 1 6, vii.26.65, P. J. Spangler (USNM).

PANAMA. — Algarrobos, 9 m. W., 8 6, 9 2, vii.5.67; Gamboa, 4 m. W.,
44,8 4, vii.6.67, P. J. Spangler (USNM); Jarones, 1 6, 2 9, vii.5.67; Las
Lajas, 4.8 m. NW., 2 6, 2 9, vil.5.67, P. J. Spangler (USNM).

MEXICO. — CHIAPAS. Huixtla, 5 m. NW., 10 6, 9 9, viti.22.65, P. J.
Spangler (USNM). TABASCO. Villahermosa, 5 m. S., 6 6, 4 2, xi.26.63,
JRZ (NMSU). VERACRUZ. J. D. Covarrubia, 6 2, vii.26.62; Paso del
Toro, 1 4, viii.25.62, JRZ (NMSU).

LITERATURE CITED

ANDERSON, R. D. 1962. The Dytiscidae (Coleoptera) of Utah; keys, origi-
nal citation, types and Utah distribution. Great Basin Nat., 22:54-75.
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Vol. 6, Méquigon, Paris, xvi + 804 pp.

JAMES R. ZIMMERMAN 235

BALFOUR-BROWNE, F. 1932. A textbook of practical entomology. Long-
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1940. British water beetles. Vol. 1. Ray Society, London,
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BALFOUR-BROWNE, J. 1944. New names and new synonymies in the Dyti-
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BLACKWELDER, R.E. 1939. Coleoptera of America, north of Mexico. Fourth
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1944. Checklist of the coleopterous insects of Mexico, Central
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, AND R. M. BLACKWELDER. 1948. Fifth supplement 1939 to
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BLATCHLEY, W. S. 1910. An illustrated descriptive catalogue of the Cole-
optera or beetles known to occur in Indiana. Indiana Dept. Geol. Nat.
Res. Bull. 1, 1386 pp.

1919. Insects of Florida. Va. Supplementary notes on the
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Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis.
Univ. California Publ. Ent., vol. 37, pp. iv + 214.

CrotcH,G.R. 1873. Revision of the Dytiscidae of the United States. Trans.
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DARLINGTON, P. J., JR. 1936. The cenotes of Yucatan; a zoological and
hydrographic survey. XVI. Aquatic Coleoptera from Yucatan. Carnegie
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Fatt, H.C. 1917. New Dytiscidae. Jour. New York Ent. Soc., 25:163-182.
GERMAR, E. F. 1824. Insectorum species novae aut minus cognitae, descrip-
tionibus illustratae. Coleoptera. Halae (Hendel), pp. xxiv + 624.
Gorpon, R. D., AND R. L. Post. 1965. North Dakota water beetles. North
Dakota insects, publ. 5, Dept. Ent., North Dakota St. Univ., pp. 1-53.
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+ 340.

Horn, G. H. 1871. Descriptions of new Coleoptera of the United States,
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1883. Miscellaneous notes and short studies of North Ameri-
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MEM. AMER. ENT. SOC., 26

236 THE GENUS LACCOPHILUS

La Rivers, I. 1951. Nevada Dytiscidae (Coleoptera). Amer. Midl. Nat.,
45 (2) :392-406.

LeacH, W. E. 1917. The zoological miscellany, 3:151.

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JAMES R. ZIMMERMAN 237

Say, T. 1823. Descriptions of insects of the families of Carabici and Hydro-
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1882b. Tribe Adephaga (continued). Families Haliplidae,
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and Dytiscidae of Ontario and Quebec. Canadian Ent., 27(3) :69-76.
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or protective coloration. Ann. Ent. Soc. Amer., 53(3):422-425.

MEM. AMER. ENT. SOC., 26

238 THE GENUS LACCOPHILUS

YounG, F. N. 1963. A new species Laccophilus from the Bahamas (Coleop-
tera: Dytiscidae). Amer. Mus. Nov., (2152) :1-5.

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141-152.

, AND H. C. SEVERIN. 1957. Distribution of the genus Laccophi-
lus in South Dakota. Jour. Kansas Ent. Soc., 30(1) :29-32.

ZIMMERMANN, A. 1920. Dytiscidae, Haliplidae, Hydrobiidae, Amphizoidae.

In Junk and Schenkling, Coleopterum Catalogus, Pars 71, 326 pp.

JAMES R. ZIMMERMAN 239

Figures 28-38. Laccophilus maculosus: 28, ventral view of male metaster-
num, metacoxal file, and postcoxal processes; 29, prosternal process; 30, male
last ventral segment; 31, female last ventral segment; 32, male oval plate; 33,
lateral view of aedeagus and parameres of L. m. maculosus; 34, lateral view of
aedeagus and parameres of L. m. decipiens; 35, lateral view of distal half of
aedeagus of L. m. shermani; 36, dorso-lateral view of distal half of aedeagus
of L. m. shermani; 37, dorso-lateral view of aedeagus of L. m. decipiens; 38,
dorso-lateral view of aedeagus and parameres of L. m. maculosus.

MEM. AMER. ENT. SOC., 26

240 THE GENUS LACCOPHILUS

Figures 39-53. Laccophilus fasciatus: 39, ventral view of male metaster-
num, metacoxal file, and postcoxal processes; 40, prosternal process; 41, male
last ventral segment; 42, lateral view of ovipositor; 43, female last ventral seg-
ment; 44, male oval plate; 45, right lateral view of distal half of aedeagus of
L. f. rufus; 46, right lateral view of aedeagus and parameres of L. f. terminalis;
47, right lateral view of distal half of aedeagus of L. f. fasciatus; 48, dorsal view
of aedeagus and parameres of L. f. terminalis; 49, dorsal view of distal half of
aedeagus of L. f. rufus; 50, dorsal view of distal half of aedeagus of L. f. fasci-
atus; 51, left lateral view of aedeagus of L. f. fasciatus; 52, left lateral view of
distal half of aedeagus of L. f. rufus; 53, left lateral view of aedeagus and left
paramere of L. f. terminalis.

JAMES R. ZIMMERMAN 241

Figures 54-61. Laccophilus proximus: 54, ventral view of male metaster-
num, metacoxal file, and postcoxal processes; 55, prosternal process; 56, male
last ventral segment; 57, female last ventral segment; 58, male oval plate; 59-61,
right lateral, dorsal, and left lateral view of aedeagus and parameres.

MEM. AMER. ENT. SOC., 26

nN
aS
iw)

THE GENUS LACCOPHILUS

64

lly
, fi Muy,

66 65

6/ 68 (SS)

Figures 62-69. Laccophilus salvini: 62, ventral view of male metasternum,
metacoxal file, and postcoxal processes; 63, prosternal process; 64, male last
ventral segment; 65, female last ventral segment; 66, male oval plate; 67-69,
right lateral view, dorsal view, and left lateral view of aedeagus and parameres.

JAMES R. ZIMMERMAN 243

Figures 70-81. Laccophilus mexicanus: 70, ventral view of metasternum,
metacoxal file, and postcoxal processes; 71, prosternal process; 72, male last
ventral segment of L. m. mexicanus; 73, female last ventral segment of L. m.
mexicanus; 74, female last ventral segment of L. m. oaxacensis; 75, male oval
plate, 76-78, left lateral views of the distal half of aedeagi of L. m. mexicanus,
L. m. atristernalis, L. m. oaxacensis; 79-81, dorsal view of aedeagi and para-
meres of L. m. oaxacensis, L. m. atristernalis, and L. m. mexicanus.

MEM. AMER. ENT. SOC., 26

244 THE GENUS LACCOPHILUS

Figures 82-89. Laccophilus pseudomexicanus: 82, ventral view of meta-
sternum, metacoxal file, and postcoxal processes; 83, prosternal process; 84,
male last ventral segment; 85, female last ventral segment; 86, male oval plate,
87-89, dorsal, right lateral, and left lateral views of aedeagus and parameres.

JAMES R. ZIMMERMAN 245

we WO

ip
Y/ yp.
1 / f/f Ih Uy GY
4f/ /

Figures 90-97. Laccophilus mistecus: 90, ventral view of metasternum,
metacoxal file, and postcoxal processes; 91, prosternal process; 92, male last
ventral segment; 93, female last ventral segment; 94, oval plate; 95, c. x. through
oval plate showing ventral crest; 96-97, left lateral and dorsal views of aedeagus
and parameres.

MEM. AMER. ENT. SOC., 26

246 THE GENUS LACCOPHILUS

MM My,
Y

Figures 98-105. Laccophilus spergatus: 98, ventral view of metasternum,
metacoxal file, and postcoxal processes; 99, prosternal process; 100, male last
ventral segment; 101, female last ventral segment; 102, male oval plate; 103,
c. x. through oval plate showing ventral crest and raised lines; 104-105, right
lateral and dorsal views of aedeagus and parameres.

JAMES R. ZIMMERMAN 247

Figures 106-113. Laccophilus fuscipennis: 106, ventral view of metaster-
num, metacoxal file, and postcoxal processes; 107, prosternal process; 108, male
last ventral segment; 109, female last ventral segment; 110, male oval plate,
111-113, right lateral, dorsal, and left lateral views of aedeagus and parameres.

MEM. AMER. ENT. SOC., 26

248 THE GENUS LACCOPHILUS

Figures 114-121. Laccophilus youngi: 114, ventral view of metasternum,
metacoxal file, and postcoxal processes; 115, prosternal process; 116, male last
ventral segment; 117, female last ventral segment; 118, male oval plate; 119-
121, right lateral, dorsal, and left lateral views of aedeagus and parameres.

JAMES R. ZIMMERMAN 249

ae

Figures 122-138. Laccophilus pictus: 122, ventral view of metasternum,
metacoxal file, and postcoxal processes; 123, prosternal process; 124-125, male
last ventral segments of L. p. pictus and L. p. insignis; 126, female last ventral
segment; 127-128, male oval plates of L. p. pictus and L. p. insignis; 129-131,
right lateral views of aedeagus and parameres of L. p. coccinelloides and aedeagi
of L. p. pictus and L. p. insignis; 132-134, dorsal views of distal half of aedeagi
of L. p. coccinelloides, L. p. pictus, and L. p. insignis; 135-138, left lateral
views of aedeagi of L. p. insignis, L. p. coccinelloides, and intergrade of L. p.
pictus and L. p. coccinelloides, and L. p. pictus.

MEM. AMER. ENT. SOC., 26

250 THE GENUS LACCOPHILUS

Figures 139-154. Laccophilus vacaensis: 139, ventral view of metasternum,
and postcoxal processes; 140, prosternal processes; 141, last two ventral seg-
ments of male; 142, female last ventral segment; 143, male oval plate; 144,
lateral view of male protarsus; 145-147, dorsal views of aedeagi and parameres
of L. v. vacaensis from Kleberg Co., Texas, L. v. chihuahuae, and L. v. vaca-
ensis from Progreso, Yucatan; 148-150, right lateral views of aedeagi and para-
meres of L. v. vacaensis from Kleberg Co., Texas, L. v. chihuahuae, and L. v.
vacaensis from Progreso, Yucatan; 151-154, left lateral views of aedeagi and
parameres of L. v. vacaensis from Kleberg Co., Texas, L. v. chihuahuae, and
L. v. vacaensis from Progreso, Yucatan, and L. v. thermophilus from La Paz,
Baja California.

JAMES R. ZIMMERMAN 251

SiN) Nae

155

158

Figures 155-162. Laccophilus spangleri: 155, ventral view of metasternum
and postcoxal processes; 156, prosternal process; 157, male last ventral segment;
158, female last ventral segment; 159, male oval plate; 160-162, right lateral,
dorsal, and left lateral views of aedeagi and parameres.

MEM. AMER. ENT. SOC., 26

252 THE GENUS LACCOPHILUS

ea

163

a

Figures 163-172. Laccophilus peregrinus: 163, ventral view of metaster-
num and postcoxal processes; 164, prosternal process; 165, male last ventral
segment; 166, female last ventral segment; 167, male oval plate; 168, lateral view
of male protarsus; 169-171, right lateral, dorsal, and left lateral view of aede-
agus and parameres; 172, lateral view showing weak line above the epipleuron.

JAMES R. ZIMMERMAN 253

175

173

178

ars

182

Figures 173-182. Laccophilus raitti: 173, ventral view of metasternum and
postcoxal processes; 174, prosternal process; 175, male last ventral segment;
176, female last ventral segment; 177, male oval plate; 178, lateral view of male
protarsus; 179, lateral view showing strong crease above the epipleuron; 180-
182, right lateral, dorsal, and left lateral views of aedeagus and parameres.

MEM. AMER. ENT. SOC., 26

THE GENUS LACCOPHILUS

nN
nN
-

185

I89

Figures 183-191. Laccophilus huastecus: 183, ventral view of metasternum
and postcoxal processes; 184, prosternal process; 185, male last ventral segment;
186, female last ventral segment; 187, distal part of male oval plate; 188, ovi-
positor; 189-191, right lateral, dorsal, and left lateral views of aedeagus and
parameres.

JAMES R. ZIMMERMAN 255

Figures 192-200. Figs. 192-198. Laccophilus quadrilineatus: 192, ventral
view of metasternum and postcoxal processes; 193, prosternal process; 194, male
last ventral segment; 195, female last ventral segment; 196, ovipositor; 197, dis-
tal half of male oval plate; 198, left lateral view of the distal third of aedeagus.
Figs. 199-200. Laccophilus sonorensis: left and right lateral views of aedeagus
and parameres.

MEM. AMER. ENT. SOC., 26

tN
Nn
On

THE GENUS LACCOPHILUS

Figures 201-208. Laccophilus biguttatus: 201, ventral view of metaster-
num and postcoxal processes; 202, prosternal process; 203, male last ventral
segment; 204, female last ventral segment; 205, male oval plate; 206-208, right
lateral, dorsal, and left lateral views of aedeagus and parameres.

JAMES R. ZIMMERMAN 257

2 II

214 AS 216

(a

Figures 209-216. Laccophilus horni: 209, ventral view of metasternum
and postcoxal processes; 210, prosternal process; 211, male last ventral segment;
212, female last ventral segment; 213, male oval plate; 214-216, right lateral,
dorsal, and left lateral views of aedeagus and parameres.

MEM. AMER. ENT. SOC., 26

258 THE GENUS LACCOPHILUS

225

222 223

Figures 217-225. Laccophilus leechi: 217, ventral view of metasternum
and postcoxal processes; 218, prosternal process; 219, male last ventral segment;
220, female last ventral segment; 221, male oval plate; 222-224, right lateral,
dorsal, left lateral views of aedeagus and parameres; 225, ovipositor.

JAMES R. ZIMMERMAN 259

ZS

230

23

Figures 226-233. Laccophilus undatus: 226, ventral view of metasternum
and postcoxal processes; 217, prosternal process; 228, male last ventral segment;
229, female last ventral segment; 230, male oval plate; 231-233, right lateral,
dorsal, and left lateral views of aedeagus and parameres.

MEM. AMER. ENT. SOC., 26

260 THE GENUS LACCOPHILUS

258

AXE |

Figures 234-241. Laccophilus schwarzi: 234, ventral view of metasternum
and postcoxal processes; 235, prosternal process; 236, male last ventral segment;
237, female last ventral segment; 238, oval plate; 239-241, right lateral, dorsal,
and left lateral views of aedeagus and parameres.

.1 JAMES R. ZIMMERMAN 261

246

Ya
| ly

Figures 242-249. Laccophilus duplex: 242, ventral view of metasternum
and postcoxal processes; 243, prosternal process; 244, male last ventral segment;
245, female last ventral segment; 246, male oval plate; 247-249, right lateral,
dorsal, and left lateral views of aedeagus and parameres.

\

MEM. AMER. ENT. SOC., 26

262 THE GENUS LACCOPHILUS

250

259 256 Zo

Figures 250-259. Laccophilus oscillator: 250, ventral view of metasternum
and postcoxal processes; 251, prosternal process; 252, male last ventral segment;
253, female last ventral segment; 254, male oval plate; 255-257, right lateral,
dorsal, and left lateral views of aedeagus and parameres of L. o. oscillator; 258-
259, right and left lateral views of the distal half of aedeagus of L. o. laevipennis.

JAMES R. ZIMMERMAN 263

265 266 267

Y

Figures 260-267. Laccophilus gentilis: 260, ventral view of metasternum
and postcoxal processes of L. g. suavis; 261, prosternal process; 262, male last
ventral segment; 263, female last ventral segment; 264, male oval plate; 265-
267, right lateral, dorsal, and left lateral views of aedeagus and parameres.

MEM. AMER. ENT. SOC., 26

264 THE GENUS LACCOPHILUS

270

AU®

Figures 268-275. Laccophilus ovatus zapotecus: 268, ventral view of meta-
sternum and postcoxal processes; 269, prosternal process; 270, male last ventral
segment; 271, female last ventral segment; 272, male oval plate; 273-275, right
lateral, dorsal, and left lateral views of aedeagus and left parameres.

JAMES R. ZIMMERMAN 265

MM

Figures 276-281. Fig. 276. Laccophilus maculosus shermani, female, Co-
chise Co., Arizona, with large epipleural flange. Fig. 277. Laccophilus macu-
losus shermani, female, Lincoln Co., New Mexico, with small epipleural flange.
Fig. 278. Laccophilus maculosus decipiens, female, San Diego Co., California,
with large epipleural flange. Fig. 279. Laccophilus maculosus decipiens, fe-
male, Alberta, Canada. Fig. 280. Intergrade of L. m. maculosus and L.-m.
decipiens, female, Sedgwick Co., Colorado. Fig. 281. Laccophilus maculosus
maculosus, male, Monroe Co., Indiana.

MEM. AMER. ENT. SOC., 26

266 THE GENUS LACCOPHILUS

286 287 MM 288

Figures 282-288. Fig. 282. Laccophilus fasciatus terminalis, female, Hi-
dalgo Co., New Mexico. Fig. 283. Laccophilus fasciatus terminalis, male, Rio
Nazas, Durango, Mexico. Fig. 284. Laccophilus fasciatus rufus, male, Mon-
roe Co., Indiana. Fig. 285. Laccophilus fasciatus rufus, female, Monroe Co.,
Indiana. Fig. 286. Intergrade of L. f. rufus and L. f. terminalis, Sedgwick
Co., Kansas. Fig. 287. Intergrade of L. f. fasciatus and L. f. terminalis, male,
Elota, Sinaloa, Mexico. Fig. 288. Laccophilus fasciatus fasciatus female, Juan
Dias Covarrubia, Veracruz, Mexico.

JAMES R. ZIMMERMAN 267

(23) ©) (35)

Figures 289-295. Fig. 289. Laccophilus proximus, female, Monroe Co.,
Indiana. Fig. 290. Laccophilus fuscipennis, male, Mazatlan, Sinaloa, Mexico.
Fig. 291. Laccophilus salvini, male, Ixtapa, Chiapas, Mexico. Fig. 292. Lac-
cophilus salvini, female, Ixtapa, Chiapas, Mexico. Fig. 293. Laccophilus
mexicanus atristernalis, male, Lincoln Co., Nevada. Fig. 294. Laccophilus
mexicanus mexicanus, male, San Juan del Rio, Durango, Mexico. Fig. 295.
Laccophilus mexicanus oaxacensis, male, El Tule, Oaxaca, Mexico.

MEM. AMER. ENT. SOC., 26

268 THE GENUS LACCOPHILUS

298

Figures 296-299. Fig. 296. Laccophilus pseudomexicanus, male, Maza-
mitla, Jalisco, Mexico. Fig. 297. Laccophilus spergatus, female, Navios,
Durango. Fig. 298. Laccophilus mistecus mistecus, male, Ixtlan de Juarez,
Oaxaca, Mexico. Fig. 299. Laccophilus mistecus aztecus, male, Atlocomulco,
Mexico, Mexico.

JAMES R. ZIMMERMAN 269

301 : SOZ
ZOO
303 304
MM

Figures 300-304. Fig. 300. Laccophilus pictus coccinelloides, female, J.
Davis Co., Texas. Fig. 301. Laccophilus pictus pictus, female, El Cameron,
Oaxaca, Mexico. Fig. 302. Laccophilus pictus insignis, male, Mitlapa, San
Luis Potosi, Mexico. Fig. 303. Intergrade of L. p. pictus and L. p. cocci-
nelloides, male, Autlan, Jalisco, Mexico. Fig. 304. Intergrade of L. p. pictus
and L. p. insignis, male, Papantla, Veracruz, Mexico.

MEM. AMER. ENT. SOC., 26

270 THE GENUS LACCOPHILUS

309

Figures 305-310. Fig. 305. Laccophilus youngi, male, La Huerta, Jalisco,
Mexico. Fig. 306. Laccophilus quadrilineatus quadrilineatus, female, Kiowa
Co., Colorado. Fig. 307. Laccophilus quadrilineatus quadrilineatus, male, Hi-
dalgo Co., New Mexico. Fig. 308. Laccophilus sonorensis, male, Hidalgo Co.,
New Mexico. Fig. 309. Laccophilus quadrilineatus tehuanensis, male, Tehu-
antepec, Oaxaca, Mexico. Fig. 310. Laccophilus quadrilineatus mayae, male,
Progreso, Yucatan, Mexico.

JAMES R. ZIMMERMAN 271

SI 312 SS

Figures 311-315. Fig. 311. Laccophilus raitti, male, San Blas, Nayarit,
Mexico. Fig. 312. Laccophilus vacaensis chihuahuae, female, Cochise Co.,
Arizona. Fig. 313. Laccophilus vacaensis vacaensis, female, Matamoros, Ta-
maulipas, Mexico. Fig. 314. Laccophilus vacaensis vacaensis, female, Liberia,
Costa Rica. Fig. 315. Laccophilus spangleri, male, Tehuantepec, Oaxaca,
Mexico.

MEM. AMER. ENT. SOC., 26

nN
~
N

THE GENUS LACCOPHILUS

Figures 316-323. Fig. 316. Laccophilus peregrinus peregrinus, female, San
Blas, Nayarit, Mexico. Fig. 317. Laccophilus peregrinus peregrinus, female,
Garro, Veracruz, Mexico. Fig. 318. Laccophilus peregrinus variabilis, male,
Ixtapa, Chiapas, Mexico. Fig. 319. Laccophilus peregrinus variabilis, male,
Anton, Panama. Fig. 320. Laccophilus huastecus, male, Tantoyuca, Veracruz,
Mexico. Fig. 321. Laccophilus biguttatus, male, Benson Co., North Dakota.
Fig. 322. Laccophilus undatus, female, Monroe Co., Indiana. Fig. 323. Lac-
cophilus schwarzi, female, Amherst Co., Virginia.

JAMES R. ZIMMERMAN

N
—
WwW

Figures 324-325. Fig. 324. Laccophilus horni, male, Tzitzio, Michoacan,
Mexico. Fig. 325. Laccophilus leechi, female, Concordia, Sinaloa, Mexico.

MEM. AMER. ENT. SOC., 26

274 THE GENUS LACCOPHILUS

Figures 326-332. Fig. 326. Laccophilus oscillator oscillator, female, Hua-
chinera, Sonora, Mexico. Fig. 327. Laccophilus oscillator laevipennis, female,
Chiapa de Corzo, Chiapas, Mexico. Fig. 328. Laccophilus duplex, male, Las
Cruces, Chiapas, Mexico. Fig. 329. Laccophilus ovatus zapotecus, male, Villa-
hermosa, Tabasco, Mexico. Fig. 330. Laccophilus ovatus zapotecus, male,
Huixtla, Chiapas, Mexico. Fig. 331. Laccophilus gentilis gentilis, female, High-
land Co., Florida. Fig. 332. Laccophilus gentilis suavis, male, Juan Diaz,
Covarrubia, Veracruz, Mexico.

JAMES R. ZIMMERMAN Das

INDEX

new species or subspecies in boldface, synoryms in italics

americanus, 91

apicalis, 70

biguttatus, 193
californicus, 46
championi, 229
confusus, 91

duplex, 214

fasciatus, 63

fasciatus fasciatus, 70
fasciatus rufus, 75
fasciatus terminalis, 80
fuscipennis, 136
fusculus, 46

gentilis, 225

gentilis gentilis, 226
gentilis suavis, 229
horni, 199

huastecus, 174
inconspicuus, 193
lateralis, 199

leechi, 205

maculosus, 29
maculosus decipiens, 46
maculosus maculosus, 33
maculosus shermani, 56
mexicanus, 109
mexicanus atristernalis, 119
mexicanus mexicanus, 113
mexicanus oaxacensis, 122
mistecus, 127

mistecus aztecus, 129
mistecus mistecus, 128

MEM. AMER. ENT. SOC., 26

optatus, 214

oscillator, 218

oscillator laevipennis, 223
oscillator oscillator, 222
ovatus zapotecus, 231
peregrinus, 166, 168
peregrinus peregrinus, 170
peregrinus variabilis, 172
pictus, 142

pictus coccinelloides, 150
pictus insignis, 154

pictus pictus, 146
proximus, 91
pseudomexicanus, 124
quadrilineatus, 180
quadrilineatus mayae, 186
quadrilineatus quadrilineatus, 182
quadrilineatus tehuanensis, 185
raitti, 177

salvini, 102

schwarzi, 211

sonorensis, 188

spangleri, 162

spergatus, 133

truncatus, 46

undatus, 208

vacaensis, 156

vacaensis chihuahuae, 160
vacaensis thermophilus, 162
vacaensis vacaensis, 158
youngi, 139

MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY
An irregular serial of monographic papers by students of authority in their

respective subjects.
1, The Cresson Types of Hymenoptera. Ezra T. Cresson. 1916. $6.00.

26,

. A Venational Study of the Suborder Zygoptera (Odonata), with Keys for the

Identification of Genera, Philip A. Munz. 1919. $2.00.

. The Blattidae of Panama. Morgan Hebard. 1920. $3.00.
. The Types of Hymenoptera in the Academy of Natural Sciences of Philadelphia

other than those of Ezra T. Cresson. Ezra T. Cresson. 1928. $2.00,

. Revision of the Rhipiphoridae of North and Central America (Coleoptera).

Ezekiel Rivnay. 1929. $2.00.

. A Revision of the Dipterous Family Rhagionidae (Leptidae) in the United

States and Canada. Mortimer D. Leonard. 1930. $4.50,

. The Eumastacinae of Southern Mexico and Central America. James A. G. Rehn

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. The Generic Names of the Sphecoid Wasps and their type species. V. S$. L.

Pate. 1937. $2.50.

A Revision of the North American species belonging to the genus Pegomyia,
H. C. Huckett. 1941. $3.00.

. Catalogue and reclassification of the Nearctic Ichneumonidae. Henry K.

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Caraga of North America (Microlepidoptera). Annette F, Braun. 1948.
50.

Classification of the Blattaria as Indicated by their Wings (Orthoptera). Johan

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. The Neotropical Species of the “Subgenus Aeschna” sensu Selysii 1883 (Odo-

nata). Philip P. Calvert. 1956. $10.00.

A Taxonomic Study of the North American Licinini with Notes on the Old
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A Taxonomic Study of the Milliped Family Spirobolidae (Diplopoda: Spiro-
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The Genus Bucculatrix in America North of Mexico (Microlepidoptera). Ann-
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The Butterflies of Liberia. Richard M. Fox, Arthur W. Lindsey, Jr., Harry K.
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A Revision of the Mexican and Central American Spider Wasps of the Subfamily
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. A Taxonomic and Zoogeographic Survey of the Scarabaeinae of the Antilles

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A Monograph of the Ithomiidae (Lepidoptera) Part If. The Tribe Mechanitini
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. A List of New North American Spiders, 1940-1966. Beatrice R. Vogel. 1967.

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. The Schizopteridae (Hemiptera: Heteroptera) With the Description of New

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A Taxonomic Revision of the Aquatic Beetle Genus Laccophilus (Dytiscidae)
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