Thesis B Dealy - Piezogaster Generic Revision

A REVISION OF THE GENUS PIEZOGASTER AMYOT & SERVILLE
(HETEROPTERA: COREIDAE: NEMATOPODINI)
AND THE DESCRIPTION OF
TWO NEW SPECIES
being
A Thesis Presented to the Graduate Faculty
of the Fort Hays State University in
Partial Fulfillment of the Requirements for
the Degree of Master of Science
by
Beau Dealy
B.S., Fort Hays State University
Date___________________ Approved_____________________________
Major Professor
Approved__________________________________
Chair, Graduate Council
Index
Acknowledgments..........................................................................................................iii
Abstract.........................................................................................................................1
Introduction...................................................................................................................2
Methods.........................................................................................................................4
Results...........................................................................................................................6
Description of the genus Piezogaster Amyot & Serville..................................................8
Diagnostic key to known species....................................................................................10
Species descriptions (alphabetical listing) –
Piezogaster achillelus Brailovsky & Barrera......................................................13

Piezogaster achilles (Stål)..................................................................................14
Piezogaster acuminatus Brailovsky....................................................................16
Piezogaster alienatus sp. nov..............................................................................18
Piezogaster alternatus (Say)...............................................................................20
Piezogaster auriculatus (Stål).............................................................................24
Piezogaster basilicus Brailovsky & Barrera........................................................27
Piezogaster bolivianus Brailovsky......................................................................30
Piezogaster calcarator (Fabricius)......................................................................30
Piezogaster camposi (Montandon).....................................................................33
Piezogaster chiriquinus (Distant).......................................................................35
Piezogaster chontalensis (Distant)......................................................................38
Piezogaster congruus Brailovsky & Barrera.......................................................40
Piezogaster dilatatus (Dallas).............................................................................41
Piezogaster humerosus (Distant)........................................................................43
i
Piezogaster indecorus (Walker)..........................................................................44
Piezogaster loricata (Distant).............................................................................46
Piezogaster multispinus (Stål)............................................................................47
Piezogaster obscuratus (Montandon).................................................................48
Piezogaster odiosus (Stål)..................................................................................50
Piezogaster reclusus Brailovsky & Barrera.........................................................52
Piezogaster rubronotatus (Stål)..........................................................................55
Piezogaster rubropictus (Montandon)................................................................57
Piezogaster scutellaris Stål.................................................................................59
Piezogaster spurcus (Stål)..................................................................................61
Piezogaster tetricus (Stål)...................................................................................63
Piezogaster thoracicus (Distant).........................................................................66
Piezogaster vates (Stål)......................................................................................67
Piezogaster yonkei sp. nov..................................................................................69
References......................................................................................................................72
Figures -
Distributional data (Fig. 1-31)...........................................................................76
Images of specimens (Fig. 32-121)....................................................................92
Illustrations (Fig. 122-144)..............................................................................116
Relative lengths for all species (Fig. 145).........................................................121
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Acknowledgments
I would like to thank the following individuals and organizations, for without their
generous encouragement, assistance and support, this career milestone would not have
been attainable:
Dr. Richard Packauskas, Angie Kuhn, Mike Dealy, Nolan Dealy, Holly Dealy, Juanita
Dealy, Eric Carver, Brant Kelsey and Jason Wenke; Mick Webb and The Natural History
Museum, London; Dr. Robert Davidson, John Rawlins, and the Carnegie Museum of
Natural History, Pittsburgh; Dr. Robert Brooks and the University of Kansas, Lawrence;
Dr. Robert Sites, Kris Simpson and the Wilbur R. Enns Entomology Museum, University
of Missouri, Columbia; Dr. Harry Brailovsky and the University of Mexico, Mexico City;
Dr. Joseph Thomasson, Dr. Jerry Choate, Dr. Greg Farley, and everyone else in the Fort
Hays Biology Department; Dr. John Heinrichs; Carolyn Herrman and Forsyth Library; Dr.
Toby Schuh and the American Museum of Natural History, NewYork; Dr. Norman Penny
and the California Academy of Sciences, San Francisco, E. Richard Hoebeke and Cornell
University, Ithaca; Daniel Summers and the Field Museum, Chicago; Kathleen Zeiders and
the Illinois Natural History Survey, Champaign; David Furth, Nancy Adams and the
National Museum of Natural History, Washington, DC; Dr. Joseph Schaffner, Edward
Riley and Texas A & M University, College Station; Cheryl Barr and the University of
California, Berkeley; S. L. Heydon and the University of California, Davis; Mark O'Brien
and the University of Michigan, Ann Arbor; and Philip Clausen and the University of
Minnesota, St. Paul.
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A Revision of the Genus Piezogaster Amyot & Serville
(Heteroptera: Coreidae: Nematopodini)
and the Description of
Two New Species
Abstract
The New World Heteropteran genus Piezogaster is revised with in-depth
descriptions of the genus and its contained species. A diagnostic key to the twenty-nine
currently recognized species, complete with images and illustrations is provided. Two
new species of Piezogaster are described, one exclusive to western Mexico, and the other
extending from the southwest United States through central Mexico. Piezogaster
alternatus (Say) is resurrected from synonymy under Piezogaster calcarator (Fabricius),
after examination of homoeotype specimens, ranges and examination of the descriptions of
both species in the literature. Piezogaster ashmeadi (Montandon) is synonymized with P.
alternatus, based on like diagnostic characters in the literature, sympatric ranges, and the
complete lack of identified specimens of P. ashmeadi. Piezogaster herrichi (Blöte) is
synonymized with Piezogaster indecorus (Walker), because of the vagueness of the
original description of P. herrichi, the reliance on an inconsistent diagnostic character for
separation, and sympatric ranges of both species. Piezogaster humeralis (Distant) is
synonymized with Piezogaster camposi (Montandon), based on prior research by O'Shea
(1974), similar diagnostic characters in both original descriptions, and material from
Distant's collection at the British Museum identified as P. humeralis that is identical in
1
2
every way to identified specimens of P. camposi. Piezogaster scitus Brailovsky & Barrera
is synonymized with Piezogaster auriculatus (Stål) after examination of original
descriptions and difficulty in finding any real character distinction between specimens of
both species, using specimens identified by Brailovsky.
Distributional data are updated by state for the United States and Mexico, as well as
by country for Central and South America. Ranges are expanded for nearly all examined
species. Distributions examined using Geographic Information System software clarified
questions concerning range overlap of P. alternatus and P. calcarator as well as P.
herrichi and P. indecorus. Distributional data analysis also showed a discrepancy in the
reported distribution of P. auriculatus; the modified range excludes New Mexico and
Texas.
Introduction
The family Coreidae arguably contains the least-studied species of any family of
heteropterans, and, historically, has exhibited much systematic confusion. My research
attempts to shed light on the particularly neglected genus, Piezogaster, and sets the
groundwork for further research on its members.
Piezogaster Amyot & Serville (1843) is a member of the Nematopodini, a tribe split
from the tribe Mictini by O'Shea & Shaefer (1978). O'Shea & Shaefer split the Mictini
into one Old World tribe and two New World tribes, with the Nematopodini belonging to
one of the latter. O'Shea (1980) then revised all of the genera within Nematopodini and
produced a key to distinguish the genera. Subsequently, the tribe Nematopodini was
3
differentiated from other coreids by Packauskas (1994) with the combination of sulcate
(sometimes shallowly) apically unarmed tibiae, a metathoracic scent gland with two
completely separate auricles, antenniferous tubercles occupying most of the head width,
and a tylus that is at most only vaguely projecting past the jugae.
Much of the early work on species now included in Piezogaster was done under the
currently synonomized genera Archimerus Burmeister (1835) and Capaneus Stål (1862).
Distant (1893) established the monotypic genus Ojedana, which was later synonymized
under Archimerus by Montandon (1899). The genus Piezogaster had been previously
synonymized under Archimerus Burmeister by Stål (1867). O'Shea (1980) resurrected the
genus Piezogaster and gave an explanation, since restated by Henry & Froeschner (1988),
that the genus Archimerus originally was established by Burmeister to be a replacement
for the preoccupied Pachymeria Laporte (1833). Archimerus Burmeister and
Pachymeria share the same type species, Pachymeria armata Laporte (1833), which has
been in synonymy under the genus Lycambes Stål since Lethierry & Severin (1894) in the
subfamily Meropachyinae. Because the genus no longer has a type species, this nullifies
Archimerus as a valid nematopodine genus. The next available genus is Piezogaster, so
O'Shea (1980) resurrected the genus and grouped all former Archimerus species, with the
exception of the type species, under Piezogaster. He also synonymized Capaneus Stål
(1862), citing too wide a range of variability among distinguishing characters between the
two genera. This brings us to the current concept of Piezogaster, which is the focus of
my research.
Except for Lethierry & Severin's (1894) catalog, O'Shea's (1980) generic revision,
and Brailovsky & Barrera's (1984) catalog of Mexican Piezogaster, work on Piezogaster
4
has been quite sporadic and sometimes limited. Other significant contributions include
those of Dallas (1852), who described new species of Archimerus from the British
Museum collection; Stål's description of new species from Mexico (1862) and his
extensive heteropteran catalog (1870); Distant's catalog of Central American Heteroptera
and descriptions of new species (1880-1893, 1901); Brailovsky & Barrera's new species
descriptions (1983, 1984, 2000), as well as Brailovsky's new species descriptions (1993).
My objective is to provide a species-level revision of the genus Piezogaster Amyot &
Serville, using all pertinent literature and specimens to which I had access. In this paper I
produce a key to distinguish all twenty-nine species in the genus, provide detailed
descriptions of species available for study, and summarize prior descriptions for species
that were not available to me. I also resolve various inconsistencies among species in the
genus, introduce two new species, and provide textual and graphical distributional data for
each species.
Methods
I conducted an extensive literature search on Piezogaster, and compiled a list of
described species. Over 3,500 specimens were borrowed from various museums and
sorted to species using existing descriptions and available diagnoses. After noting the
characters and distributional data for all specimens, samples of up to ten males and ten
females from each species were selected for length and width measurements. When
available, annotated specimens were used. I had access to the holotype for Piezogaster
acuminatus Brailovsky, and homoeotypes, specimens compared directly with the
holotype, for Piezogaster auriculatus (Stål), Piezogaster calcarator (Fabricius),

5
Piezogaster odiosus (Stål), Piezogaster scutellaris Stål, Piezogaster spurcus (Stål), and
Piezogaster tetricus (Stål). Three measurements were taken for each specimen: length
from the tip of tylus to the end of the abdomen, pronotal width at the widest point, and
abdominal width at the widest point (Fig. 144). Measurements were conducted with an
ocular micrometer and measured to the nearest tenth of a millimeter. Two different
databases were created for the data, one for morphological characters and another for
distributions and measurements. Distributional data were entered by state for the United
States and Mexico, and by country for Central and South America. Distributional data for
South America was scant, mainly due to small sample sizes. Data for physical
characteristics were entered into a database using DELTA v.1.01 (Dallwitz, et al.1980-
1999), and all measurement and distributional data were entered into a spreadsheet. With
these data sets, I wrote detailed descriptions of species. Species that I had no specimens
for or that were poorly represented were summarized using the available literature,
making special note of distinguishing characters. From the data collected, I assembled a
dichotomous key for all known Piezogaster species. The bulk of the character data was
from the DELTA database. The keys generated using this database served as a
foundation to add inaccessible species. This was accomplished by incorporating
diagnostic characters from existing written descriptions and observations from illustrations
and photographs.
Distributional data were assembled and associated with spatial data for the western
hemisphere using ArcView 3.1 (Environmental Systems Research Institute 1992-1998) for
the Windows operating system. A data set for each species was extracted from this
assembly showing the historic distributions for each species and incorporating newly
6
discovered distributions through my research. Distributional data are given in each
species description, as well as graphically represented (Figures 1-31). New distributions
resulting from my research are bold faced in the text with the acronym of the museum that
the specimen came from following in parentheses. Acronyms for museums are: AMNH
(American Museum of Natural History, NewYork), BMNH (British Museum of Natural
History, London), CAS (California Academy of Sciences, San Francisco), CMNH
(Carnegie Museum of Natural History, Pittsburgh), CU (Cornell University, Ithaca),
FMNH (Field Museum of Natural History, Chicago), NMNH (National Museum of
Natural History, Washington, DC), SMEK (Snow Museum of Entomology, University of
Kansas, Lawrence), SMHP (Sternberg Natural History Museum of the High Plains, Fort
Hays State University, Hays), TAMU (Texas A & M University, College Station), UCB
(University of California, Berkeley), UMAA (University of Michigan, Ann Arbor), UMC
(University of Missoiri, Colombia), UMSP (University of Minnesota, St. Paul).
For all species for which specimens were available, color images were taken of the
top and side of each sex (Figures 32-117). Macro images were captured using a Nikon
Coolpix 900 series digital camera affixed to a stationary platform.
Results
From my research, I described two new species of Piezogaster, one exclusive to
western Mexico, and the other extending from the southwest United States through
central Mexico. I also raised one species from synonymy, synonymized four other
species, updated and expanded distributional data, and produced a diagnostic
dichotomous key to species.

7
Piezogaster alternatus (Say) is resurrected from synonymy under P. calcarator after
examination of homoeotype specimens, ranges and examination of the descriptions of both
species in the literature. I synonymize Piezogaster ashmeadi (Montandon) with P.
alternatus, based on like diagnostic characters in the literature, sympatric ranges, and the
complete lack of identified specimens of P. ashmeadi. Piezogaster herrichi (Blöte) is
synonymized with Piezogaster indecorus (Walker) because of the vagueness of the
original description of P. herrichi, the reliance on an inconsistent diagnostic character for
separation, and sympatric ranges of both species. Piezogaster humeralis (Distant) is
synonymized with Piezogaster camposi (Montandon) based on prior research by O'Shea
(1974), similar diagnostic characters in both original descriptions, and material from
Distant's collection at the British Museum identified as P. humeralis that is identical in
every way to identified specimens of P. camposi. Piezogaster scitus Brailovsky & Barrera
is synonymized with P. auriculatus after examination of original descriptions and my
difficulty in finding any real character distinction between specimens of both species, using
specimens identified by Brailovsky.
Distributional data are updated by state for the United States and Mexico, as well as
by country for Central and South America. Ranges are expanded for nearly all examined
species. Distributions examined using Geographic Information System software clarified
questions concerning range overlap of P. alternatus and P. calcarator as well as P.
herrichi and P. indecorus. Distributional data analysis also showed a discrepancy in the
reported distribution of P. auriculatus; the modified range excludes New Mexico and
Texas.
My research provides a centralized and up-to-date resource for Piezogaster, with the
8
first complete key to species for the genus. Distributional data provided here should set
the groundwork for a more detailed biogeographical study, and hopefully, my dealings
with the nomenclatorial problems in this genus will stimulate examination of Piezogaster
by other heteropterists, and further overall coreid taxonomic endeavours.
Genus Piezogaster Amyot & Serville
Piezogaster Amyot & Serville. 1843. p. 197. Type species: P. albonotatus Amyot &
Serville 1843. Monotypic.
Capaneus Stål. 1862. p. 279. Synonymized by O'Shea (1980). Type species: Capaneus
multispinus Stål 1862.
Archimerus: Stål (not Burmeister), 1867. p. 538.
Piezogaster: Stål, 1867.
Ojedana Distant, 1893. p. 355. Type species Ojedana loricata Distant 1893 (in Distant
1880-1893). Synonomized under Archimerus by Montandon (1899).
Piezogaster: O'Shea, 1980. Resurrected from synonymy.
Piezogaster: Baranowski & Slater, 1986. p. 38.
Piezogaster: Henry & Froeschner, 1988. p. 87.
Color variable, ranging from pale to dark brown or black to reddish or reddish-
orange. Body depressed, head somewhat quadrate, tapered at anterior end, ranging from
tubercled to tubercles vaguely present. Tylus extending well past strongly deflexed jugae;
jugae not visible dorsally. Antenniferous tubercles separated by tylus, distance subequal
to width of one antenniferous tubercle. Post ocular tubercles usually present. Antennal
segment length ratios variable, but segment III always shortest; segment I robust. Beak
always extending past prothorax, segment III always shortest. Pronotal collar always
present. Pronotum variable, especially lateral angles, but always steeply declevent (more
than 45 degrees from horizontal). Scutellum usually punctate. Lacking raised

9
mesosternal sulcus associated with the genus Mozena. Thoracic pleura punctate, often
tuberculate. Metathoracic scent gland placed laterally on pleura with both anterior and
posterior auricles, anterior auricle usually larger of the two. Each connexival segment
usually armed to some degree with at least a small spur at the lateral posterior angle. All
femora armed with at least one pair of ventrodistal spines, usually preceded by much
smaller, less conspicuous spines. Spine pattern usually consistent for all femora, but
amplified in degree and size on metafemora; male metafemora are nearly always
incrassate. Pro- and mesotibiae apparently triquetrous in cross section, male metatibiae
usually laterally compressed as well. Prominent external groove usually visible, running
length of all tibiae; males with lone medial ventral spine present. Tarsi nearly always with
fringe of small, sometimes thick, hairs surrounding distal end. Tarsi slightly less than half
the length of their associated tibia. Abdomen varies from much wider than pronotum to
much narrower than connexiva.
Measurements – Body length ranging from 29 mm to15 mm (see Fig. 145 for relative
lengths of all species); pronotal width ranging from 13 mm to 5 mm; abdominal width
ranging from 14 mm to 4 mm.
Diagnosis – A few distinct characters separate Piezogaster from other Nematopodini.
Most notable of these are a steeply declivent (at least 45 degrees from horizontal)
pronotum, a tylus that extends past the antenniferous tubercles, and a dorsal metafemoral
surface that is usually armed with tubercles or spines, which are especially prevalent in
males. Of the Nematopodini, only Piezogaster and Mozena bear these traits; however,
Piezogaster lacks the raised mesosternal medial sulcus found in Mozena in which the beak
sometimes rests.
10
Diagnostic Key to known species of the genus Piezogaster
1 Pronotal expansions present (Fig.126-135) or humeral angles markedly

widened (Fig. 140)...........................................................................................2
1' Pronotal expansions lacking, pronotal angles not widened (Fig.136-139).........17
2(1) Pronotal margin strongly dentate-serrate, femora extremely multispinose (Fig.

123).....................................................................................P. multispinus (Stål)*
2' Pronotal margin not strongly dentate-serrate; may be tuberculate or weakly
serrate (Fig. 126-140). If femoral spines present, not abundant.......................3
3(2) Pronotum covered in strong, rounded, reddish-brown to black tubercles, more

numerous on rising lateral expansions (Fig. 131) ..............P. humerosus (Distant)
3' Pronotum lacking strong, rounded, reddish-brown to black tubercles, or if present,
lacking on expansions......................................................................................4
4(3) Pronotum expanded laterally, terminating in subacute points (Fig. 130) ...........
........................................................................................P. thoracicus (Distant)*
4' Pronotum lacking lateral expansions, or if laterally expanded, lacking subacute
points..............................................................................................................5
5(4) Laterally projecting jugal shelf present beneath base of antennal segment I.......6
5' Lacking laterally projecting jugal shelf beneath base of antennal segment I ......11
6(5) Antennal segment IV concolorous with remainder of segments........................7

6' Antennal segment IV not concolorous with remainder of segments..................10
7(6) With a pale yellow sternal fascia on each thoracic pleura (Fig. 81, 83)..............
........................................................................................................................8
7' Sternal fascia lacking.......................................................................................9
8(7) Body black; with 4 pairs of pale yellow abdominal discoid areas running .........
in series................................................................................P. loricata (Distant)*
8' Body medium to dark brown, never black; abdominal discoid areas lacking or
reduced to minute or obscure spots.............................P. obscuratus (Montandon)
9(8) Pronotal expansions rounded, earlike (Fig. 135); abdomen wider than pronotum
..............................................................................................P. auriculatus (Stål)
9' Pronotal expansions not rounded, terminating in a slightly anteriorly directed acute
angle (Fig. 129); pronotum wider than abdomen ............P. chontalensis (Distant)
11
10(6) Some metafemoral tubercles pale yellow; metafemora very incrassate for
size, even in females; metatibiae only slightly arcuate.......................................
................................................................................................P. scutellaris (Stål)
10' Some metafemoral tubercles same color as lighter regions of metafemora, but
never pale yellow; metafemora not always incrassate, male metatibiae
strongly arcuate or sinuate................................................P. chiriquinus (Distant)
11(5) Lateral margins of pronotum strongly tuberculate or spinose (Fig. 36, 72, 132)
........................................................................................................................12
11' Lateral margins of pronotum at most weakly tuberculate.................................14
12(11) Ultimate antennal segment bicolored..............................P. acuminatus Brailovsky

12' Ultimate antennal segment unicolorous............................................................13
13(12) Pronotal expansions slightly pointing anteriorly......................P. dilatatus (Dallas)

13' Pronotal expansions not anteriorly pointing, but laterally produced as to form a
thinly acute lateral spine................................................P. bolivianus Brailovsky*
14(11) With dark brown to black maculae on red to reddish-orange coria...................

.......................................................................................P. camposi (Montandon)
14' Maculae lacking; overall color of coria not red to reddish-orange....................15
15(14) Pronotal expansions slightly projecting anteriorly (Fig. 127); restricted to Costa
Rica..................................................................P. reclusus Brailovsky & Barrera
15' Pronotal expansions strongly projected forward, or slightly projecting anteriorly
but not pointed anteriorly (Fig. 126, 128-129); not restricted to Costa Rica.....16
16(15) Pronotal angles extremely forward-swept (Fig. 129)........................................

.....................................................................P. achillelus Brailovsky & Barrera*
16' Pronotal angles forward-swept, but not extremely so (Fig. 128). .....................
..................................................................................................P. achilles (Stål)
17(1) Having an ocherous-orange medial longitudinal stripe extending from the head
through the scutellum; male with finger-like projection arising at end of genital
capsule (Fig. 141)...........................................P. congruus Brailovsky & Barrera*
17' Lacking ocherous-orange medial longitudinal stripe extending from the head
through the scutellum; male lacking finger-like projection arising at end of
genital capsule.................................................................................................18
18(17) Antennal segment IV concolorous with remainder of segments........................19

18' Antennal segment IV not concolorous with remainder of segments..................23
19(18) Overall body color black or nearly black in color; may have colored accents....20
19' Overall body color not black, or if black, lacking colored accents....................21
12
20(19) Pronotum margined anteriorly with red semilunar band (Fig. 94, 96)................
............................................................................................P. rubronotatus (Stål)
20' Lacking red semilunar band on pronotal margin, with four reddish-orange to
orange longitudinal stripes (Fig. 98, 100)....................P. rubropictus (Montandon)
21(19) Overall body color rose or sanguine...........................................P. yonkei sp. nov.

21' Overall body color not rose or sanguine...........................................................22
22(21) Posterior pronotal angles present; body not at all fuscous.................................

......................................................................................P. calcarator (Fabricius)
22' Posterior pronotal angles lacking; body fuscous with white hairs. ....................
....................................................................................................P. spurcus (Stål)
23(18) Posterior pronotal angles present (Fig. 135-137).............................................24

23' Posterior pronotal angles lacking (Fig. 138-140)..............................................26
24(23) Head, pronotum, femora covered with white and scattered black erect hairs;
found in North America East of the continental divide..............P. alternatus (Say)
24' Covering of erect hairs on body lacking or sparse; usually found in the American
Southwest or Mexico..........................................................P. indecorus (Walker)
25(23) Connexival segments bicolored, lighter color easily contrasting with darker
color that is easily seen with naked eye............................................................26
25' Connexival segments unicolorous, or rarely with slightly lighter areas seen only
with magnification...........................................................................................27
26(25) Of great length, 24 to 25 mm..........................P. basilicus Brailovsky & Barrera‡

26' Of medium length, 16 to 21 mm...................................................P. tetricus (Stål)
27(25) Abdomen wider than pronotum (Fig. 143) .............................P. alienatus sp. nov.
27' Abdomen narrower or nearly as wide as pronotum (Fig. 142) .........................28
28(27) Always very dark brown to black, length greater than 23 mm..........P. vates (Stål)
28' Color varies from medium brown to almost black; never longer than 23 mm....
....................................................................................................P. odiosus (Stål)
‡ Key couplet taken directly from Brailovsky & Barrera (1984). My measurements of P.basilicus and P.
tetricus fall between these measurements, but not enough material was available to confirm this as
something consistent. See P. basilicus or P. tetricus notes for details.
* Denotes unexamined species.

13
Piezogaster achillelus Brailovsky & Barrera
Piezogaster achillelus Brailovsky & Barrera. 2000; p. 275; Mexico.
This recently named species was described by Brailovsky & Barrera (2000) as
being very similar to P. achilles, "agreeing in almost all details of shape and color." They
point out that P. achillelus has pronotal expansions that are much more forward swept
(Fig. 126) than those of P. achilles, which tend to be expanded as much anteriorly as
laterally (Fig. 128). Also mentioned is the smaller size and thinner stature of P. achillelus
when compared to P. achilles. Brailovsky & Barrera also compared P. achillelus to P.
chontalensis, and remarked that both are similar with respect to size and coloration, but
that the pronotal angles of the two differ. I observed that this is probably also the case
with P. reclusus. The pronotal angles of P. chontalensis and P. reclusus are shorter and
are directed slightly posteriorly at the apex (Fig.129), and turned only slightly forward
(Fig. 127), respectively, whereas the expansions of P. achillelus are longer and and are
forward swept (Fig. 126). Brailovsky & Barrera also mention that the genital capsule of
P. achillelus has a "strong 'y' longitudinal expansion" in caudal view, which appears to be
similar to the raised, somewhat flattened, inverted trigonal area on the genital capsule of
P. achilles when viewed posteriorly. I have not had opportunity to examine specimens.
Measurements – From Brailovsky & Barrera (2000). Males: n = 35; females: n = 24.
Length (mm) – Range: 16.0 – 20.6.
Distribution – (Fig. 1) Mexico: Guerrero, Oaxaca.

14
Piezogaster achilles (Stål)

(Fig. 32-35)
Capaneus achilles Stål 1862; p. 280; Mexico.

Piezogaster achilles: O'Shea 1980; p. 214.
Body color yellowish to orangish-brown, covered with minute, almost velvety, hairs
on pleura, venter of thorax, tubercles sparse to lacking. Postocular area never darker than
remainder of head, slightly tuberculate. Antennal segment I longest; II, IV subequal, III
shortest; all segments concolorous; segment II, III hairs in parallel rows running
lengthwise. Laterally projecting jugal shelf beneath base of antennal segment I lacking.
Beak segment IV longest, I, II subequal, segment III shortest. Pronotal collar impunctate,
rarely tuberculate. Pronotum margined with sparse tubercles, surface heavily punctate.
Callar region lacking tubercles or punctation. Pronotum with prominent lateral
expansions pointing anterolaterally. Posterior pronotal angles lacking. Scutellum
punctate except at anterolateral angles. Corium concolorous with body; membrane
darker, venation of membrane rarely anastomosing. Pronotum wider than abdomen in
males; variable in females. Connexiva lacking tubercles, unicolorous. Minute
posterolateral spur present on connexival segments VI, VII, sometimes V. Abdomen
lacking ventral markings. Genital capsule rim straight, entire, with sparse hairs. Genital
capsule in posterior view rugose, covered with punctations; with a raised, somewhat
flattened, inverted trigonal area on the genital capsule when viewed posteriorly.
Propleural acetabular suture not keeled. Posterior metapleural margin straight. Area
surrounding abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob
lacking. Profemora, mesofemora with one to two pairs of slight to vague tubercles on
15
distal ventral side. Protibiae, mesotibiae vaguely triquetrous in cross-section, nearly
round. External groove running length of protibiae and mesotibiae only vaguely present.
Distal end of tibiae lacking fringe of hairs. Three to four pairs of darkened spines running
ventral length of metafemora, becoming less acute distally; spine pairs more prominent in
males. Male metatibiae arcuate, anterior face of metatibiae concolorous with posterior
face; external apical tubercle lacking.
Measurements – Males: n = 2; Female: n = 1.
Length (mm) – Males: 20.5 and 21.1, Female: 19.6.
Width at pronotum (mm) – Males: 8.7 and 9.5, Female: 8.5.
Width at connexiva (mm) – Males: 6.0 and 6.4, Female: 7.0.
Diagnosis – P. achilles may be confused with P. achillelus, P. chontalensis and P.
reclusus, all of which are lacking tubercles, have a pronotum wider than the abdomen, and
have anteriorly pointing pronotal expansions, but P. achilles usually is larger than the
other three, and there are slight differences in the shape of the pronotum. P. achilles has
expansions that extend laterally as much as they do forward (Fig. 128). P. achillelus has
pronotal expansions that are longer and much more forward swept (Fig. 126). The
expansions of P. chontalensis are directed slightly posteriorly at the apex (Fig.129) and
the expansions of P. reclusus are similar to to those of P. chontalensis but are turned
slightly forward (Fig. 127). Male P. achilles also have a raised, somewhat flattened,
inverted trigonal area on the genital capsule when viewed posteriorly, which separates it
from P. chontalensis and P. reclusus. However, Brailovsky & Barrera state that the
genital capsule of P. achillelus has a "strong 'y' longitudinal expansion" in caudal view,
which appears to be similar to what I observed on the genital capsule of P. achilles.

16
Moreover, P. achilles may be separated from P. chontalensis by the darker chestnut
coloration on the femora and pronotum of P. chontalensis.
Distribution – (Fig. 2) Costa Rica (NMNH); Guatemala; Mexico:Veracruz, San Luis
Potosi.
Piezogaster acuminatus Brailovsky

(Fig. 36-37)
(Redescription from holotype)
Piezogaster acuminatus Brailovsky 1993; p.111; Brazil.
Body color brown to ruddy brown, covered with minute white hairs on head,
pronotum, and often metafemora. Entire head and venter of body covered with prominent
tubercles. Postocular area heavily tuberculate, to the point of not exposing the surface
underneath. Antennal segments I, II, IV subequal, segment III shortest; ultimate antennal
segment bicolored, apex pale yellow to yellowish-brown; proximal end concolorous with
body. Lacking laterally projecting jugal shelf beneath base of antennal segment I. Beak
segments I, II, IV subequal, segment III shortest. Anterior pronotal collar with prominent
tubercles and punctations. Pronotum margined anteriorly with dentate tubercles; callar
area separated into two distict tuberculate regions covered in white hairs; expansions
present, extending laterally and terminating in a point. Posterior pronotal angles present.
Scutellar surface rugosely punctate. Corium same color or slightly paler than remainder
of body; membrane paler in color than remainder of body, sometimes apparently a
yellowish-tan; veins of membrane anastomosing. Abdomen wider than pronotum.
Connexiva unicolorous, with minute hairs, lacking tubercles. Minute posterolateral spur
17
present on connexival segments II through VII. Pair of small, round, yellowish fascia
present medially on the venter of abdominal segment V; segment IV also having 2 or 3
pairs of small, raised, yellowish spots medially. Genital capsule rim with two dorsal teeth;
rim margined with hairs. Genital capsule in posterior view medially rugose, punctate,
laterally minutely granular. Propleural acetabular suture not keeled. Two small tubercles
located dorsal to metathoracic scent gland; auricles yellowish, subequal in size, appearing
inflated. Posterior metapleural margin sinuous. Area surrounding abdominal segment IV
spiracle not raised as trigonal area. Metacoxal knob lacking. Pro- and mesofemora with
large ventral penultimate spine; ultimate spine minute to lacking, followed by 3-4 proximal
pairs of smaller spinose tubercles. Metafemora with similar spine arrangement, but with
ultimate, and penultimate spines fused. Proximal spinose tubercle rows obscured by
interspersed tubercles. Inner and outer metatibia faces concolorous; metatibiae nearly
straight, or vaguely arcuate. Male metatibial external apical tubercle lacking.
Measurements – Male: n = 1.
Length (mm) – 18.7.
Width at pronotum (mm) – 7.5.
Width at connexiva (mm) – 9.7.
Diagnosis – P. acuminatus is similar to P. dilatatus in overall shape, with a densely
tuberculate callar region, and a tuberculate pronotum. However, the pronotal angles of P.
acuminatus extend into a lateral spine (Fig. 36, 132), whereas the pronotal angles of P.
dilatatus curve and point anteriorly (Fig. 70, 72). Moreover, the ultimate antennal
segment of P. acuminatus is bicolored, whereas the ultimate antennal segment of P.
dilatatus is unicolorous.
18
Distribution – (Fig. 3) Brazil.
Notes – I have examined only the holotype of this species, loaned to me through the
graciousness of the Carnegie Museum of Natural History.
Piezogaster alienatus sp. nov.

(Fig. 38-41)
(Description of holotype)
Body color brownish-black to black dorsally; with reddish-orange hue ventrally.
Body covered in small white decumbent hairs, sometimes nearly velvety. Tubercles
sparse. Postocular area tuberculate, concolorous with body. Antennal segments I, II
subequal, longest; segments III, IV subequal, shorter; segment IV reddish-orange to
orange in color. Laterally projecting jugal shelf beneath base of antennal segment I
lacking. Beak segments I, II, IV subequal in length, III shortest. Lateral surfaces of
anterior pronotal collar punctate, tuberculate. Pronotal lateral expansions lacking. Callar
region impunctate, covered with velvety hairs; remainder of pronotum usually less hairy,
punctate. Posterior pronotal angles present. Scutellum rugosely punctate, angles paler in
color. Corium concolorous with body. Membrane apparently darker, nearly black;
venation with little or no anastomosis. Pronotum not wider than abdomen. Connexival
segments darkly margined and covered with minute hairs; tubercles lacking.
Posterolateral connexival spur present on segments IV, V, VI. Abdomen lacking ventral
markings. Propleural acetabular suture not keeled. Rugose area surrounding
metathoracic scent gland lighter in color, orange to orangish-yellow. Posterior
metapleural margin somewhat sinuous. Area surrounding abdominal segment IV spiracle

19
not raised as trigonal area. Pro- and mesofemora with ventrodistal pair of minute ultimate
spines; penultimate spines much larger. Metafemoral spine arrangement similar to pro-
and mesofemora, but spines fused followed by 3 pairs proximal of sometimes spinose
tubercles. Metatibial faces not bicolored.
Holotype: female (Fig. 40-41). Length: 20.9 mm; width at pronotum: 8.3 mm; width at
connexiva: 9.9 mm. Mexico: Oaxaca 2.7 mi. NW El Camaron. July 14, 1971. Collected
by Clark, Hart, Murray, and Schaffner. Holotype designated from and deposited in the
entomology collection of Texas A&M University, College Station.
Measurements of all specimens – Males: n = 12; females: n = 9.
Length (mm) – Range: 19.7 - 23.6, X = 22.0; male range: 21.9 - 23.6, X = 22.8; female
range: 19.7 - 22.0, X = 21.1.
Width at pronotum (mm) – Range: 7.9 - 9.5, X = 8.7; male range: 8.3 - 9.5, X = 8.8;
female range: 7.9 - 8.9, X = 8.4.
Width at connexiva (mm) – Range: 8.9 - 10.5, X = 9.6; male range: 8.9 - 10.1, X = 9.5;
female range: 8.9 - 10.5, X = 9.7.
Diagnosis – This species is easily separated by all others by the combination of its
brownish-black to black color, orangish-red accents on antennal segment IV and around
the metathoracic scent gland, and and abdomen wider than the pronotum (Fig. 143). The
males also possess a distinct hooked knob projecting from the metacoxa.
Distribution – (Fig. 4) Mexico: Guerrero (NMNH), Oaxaca (TAMU).
Notes – This species is named for the fact that the first examples I received were from the
Texas A&M University, which acquired the specimens from an intercepted food shipment
crossing the border to the United Stated from Mexico.

20
Male characters from paratypes. Genital capsule rim entire, apparently somewhat
dorsally convex, obscuring hairs on margin of rim. Seen from posterior view, genital
capsule with small hairs, medially punctate. Male metacoxae with prominent lateral
hooked knob; tibiae slightly arcuate with external apical tubercle lacking. Paratypes
deposited in NMNH, SMHP, and TAMU.
Piezogaster alternatus (Say)
(Fig. 42-45)
Raised from Synonomy under P. calcarator
Coreus alternatus Say 1825; p. 317; "Missouri Territory"

Archimerus squalus Klug 1835; In Burmeister p.321. Synonymized by Van Duzee under
Archimerus alternatus (1917).
Archimerus muticus Herrich-Schäffer 1842; p.52. Synonymized by Stål (1870).
Archimerus rubiginosus Herrich-Schäffer 1842; p. 83. Synonymized by Stål (1870).
Piezogaster albonotatus: Amyot & Serville 1843; p.197. Synonymized under Archimerus
calcarator by Stål (1870).
Physomerus pallens Dallas 1852; p. 412. Synonymized under Archimerus calcarator by
Stål (1870).
Archimerus alternatus: Stål 1870; p. 137.
Archimerus calcarator: Lethierry & Severin 1894; p. 17.
Archimerus ashmeadi Montandon 1899; p.194; New synonymy.
Archimerus pallens: Distant 1901; p. 416.
Piezogaster calcarator: O'Shea 1980; p. 214. In part.
Piezogaster ashmeadi: O'Shea 1980; p. 214.
Piezogaster alternatus: Baranowski & Slater 1986; p.38.
Piezogaster ashmeadi: Baranowski & Slater 1986; p.39.
Piezogaster calcarator: Henry & Froeschner 1988; p. 88. In part.
Body color medium brown. Head, pronotum, and femora covered with white and
scattered black erect hairs; head tuberculate. Postocular area darker than remainder of
head, tuberculate. Antennal segment I longest, segment II, IV subequal, segment III
shortest; segment IV often lighter in color. Laterally projecting jugal shelf beneath base of
21
antennal segment I present, but not prominent. Beak segment I longest, segments II, IV
subequal, segment III shortest. Lateral sides of pronotal collar strongly punctate.
Anterior half of pronotal margin darker, also more tuberculate than posterior half. Callar
region with tubercles, never punctate, remainder of pronotum strongly punctate. Pronotal
lateral expansions lacking. Posterior pronotal angles present. Scutellum occasionally
yellowish, usually rugosely punctate. Corium concolorous with body; membrane
apparently darker in color; venation with some anastomosis. Pronotum subequal in width
to abdomen. Connexival segments with minute hairs, tuberculate, with pale spot
anteriorly; area surrounding spot, as well as lateral margins posterior to spot margined in
dark brown or black; pale spot increasing in size as series extends posteriorly, often
becoming an entire transverse bar on segment VII. Connexival posterolateral spurs
indistinct. Pair of minute, yellowish, ventro-abdominal spots often present on segments
III-VI. Genital capsule rim entire but slightly depressed medially; margined with minute
hairs. Genital capsule in posterior view medially rugose, punctate. Propleural acetabular
suture with strong keel. Anterior metathoracic scent gland auricle sometimes yellowish in
color. Posterior metapleural margin sinuous. Area surrounding abdominal segment
spiricle IV lacking raised trigonal area. Metacoxal knob lacking. Pro- and mesofemora
with ventrodistal pair of minute ultimate spines; penultimate spines much larger, often
with 3-4 pairs of pale yellow tubercles or minute spines proximally. Metafemora with
spinose tubercles dorsally, often in lengthwise rows. Venter of metafemora with similar
arrangement to pro- and mesofemora except two most distal pairs of spines fused; spines
appearing more often than tubercles following. In males, one spine on interior of
metafemora sometimes significantly larger than remainder of metafemoral spines.

22
Anterior face of metatibiae concolorous with posterior face; male metatibiae arcuate;
external apical tubercle lacking.
Measurements – Males: n = 10; females: n = 10.

Length (mm) – Range: 17.0 - 21.0, X = 18.9; male range: 18.3 - 21.0, X=19.5; female

range: 17.0 - 20.1, X = 18.3.


female range: 6.1 - 7.4, X = 6.6.


female range: 7.2 - 8.8, X = 7.9.
Diagnosis – This species is often confused with P. indecorus in collections and in areas
where the two species exist in sympatry, sharing almost every visible chracter except the
white and scattered black erect hairs on the head, pronotum, and femora, which P.
indecorus lacks. Also, antennal segments II and III of the latter sometimes appear more
robust than those of P. indecorus, due to the covering of white and scattered black erect
hairs. P. alternatus is cosmopolitan over most of the United States east of the Continental
Divide, whereas P. indecorus is a southern species, limited to Mexico and the
southwestern United States.
Distribution – (Fig. 5) United States: Alabama (UMAA), Arkansas (UMC), Colorado,
Connecticut (NMNH), Florida, Georgia (SMEK), Illinois, Indiana (TAMU), Iowa
(TAMU), Kansas, Kentucky (NMNH), Louisiana (UMSP), Maryland (CU),
Michigan, Minnesota (SMEK), Mississippi, Missouri (UMC), New Jersey, New York
(NMNH), North Carolina, Ohio (TAMU), Oklahoma, Pennsylvania (CMNH), South
Carolina, Tennessee, Virginia (NMNH), West Virginia (NMNH), Wisconsin.

23
Notes – Apparently, O'Shea (1980) incorrectly synonymized this species with P.
calcarator. I have viewed homoeotypes of P. calcarator that were designated and labeled
as such by Thomas Yonke, which are pale brown, often with an orange hue, as opposed to
P. alternatus wich is a medium brown. The P. calcarator homoeotypes also lack the pale
yellow connexival spots, and the covering of white and scattered black erect hairs, which
are present on P. alternatus. Moreover, P. alternatus occupies much of the same range
as P. calcarator sensu Van Duzee (1909) (Fig. 31). Henry & Froeschner (1988) also
noted that O'Shea, in his synonymy of P. alternatus to P. calcarator, decided to follow
Lethierry & Severin (1894) rather than using the characters of Van Duzee (1909), who
also mentions the pale color and a lack of pale yellow connexival spots, and also
recognized that P. calcarator was found only in the southeast part of the United States.
After reviewing specimens, their distributions, and descriptions in the literature, this
appears to be an obvious case of sympatry to me. Therefore, I formally restore P.
alternatus from synonymy. Van Duzee's (1917) catalog furthers the distinction of P.
alternatus from P. calacarator, and my separations of synonyms between the two species
consequently follow his.
Furthermore, it appears that Montandon (1899) was incorrect in naming P.
ashmeadi as a species. Montandon's original description makes reference to pale yellow
connexival spots, just like P. alternatus. Unidentified specimens I have examined having
pale yellow connexival spots from Florida, Montandon's type locality for P. ashmeadi,
appear identical in all respects to those of P. alternatus from all other locations. This
combined with the comments of Baranowski & Slater (1986), as well as samples that I
received of specimens that were obviously P. alternatus and P. indecorus but which were
24
labeled as P. ashmeadi, leads me to formally synonymize P. ashmeadi with P. alternatus.
During an instance of oversight, P. albonotatus Amyot & Serville (1843) was
synonymized under Physomerus pallens Dallas (1852) by Dallas in the same publication.
Physomerus pallens then was not only synonymized under Archimerus calcarator by Stål
(1870) but also remained listed in Archimerus by Distant (1901). Apparently, Distant had
overlooked Stål's earlier work.
I have examined specimens of P. alternatus found on the following plants: Olive tree,
Helianthus sp., Desmanthium sp., Circium muticum, and Ambrosia trifidum. Yonke &
Medler (1969a) found adults feeding on Solidago altissma, Aster sagittifolius, Galium
concinnum, Erigeron annus, Symplocarpus foetidus, and Desmodium acuminatum.
Yonke & Medler also observed nymphs as well as adults feeding on various other
members of Desmodium, as well as Amphicarpa bracteata, Ambrosia artemisiifolia,
Ambrosia trifida, Cryptotaenia canadensis, and Eupatorium rugosum. Alder & Wheeler
(1984) also make reference to questionable feeding on a bird dropping, however,
Packauskas (personal communication) says this phenomenon is quite common among
coreids. Yonke & Medler (1969b) provided a rather detailed look at the immature stages,
complete with illustrations.
Piezogaster auriculatus (Stål)

(Fig. 46-49)
Capaneus auriculatus Stål 1862; p. 289; Mexico.

Xuthus auriculatus Uhler 1876; p.296.
Piezogaster auriculatus:O'Shea 1980; p. 214.
Piezogaster scitus Brailovsky & Barrera 1984; p.134. New synonymy.
25
Body color dark orange to dark reddish-orange, covered with minute hairs; body
strongly tuberculate. Postocular area darker in color than remainder of head, tuberculate.
Antennal segment I tuberculate, segments II, III sometimes tuberculate; segment I
longest, segments II, IV subequal, segment III shortest; all segments concolorous.
Laterally projecting jugal shelf present beneath base of antennal segment I. Beak
segments I, II longest, subequal; segment IV shorter, segment III shortest. Anterior
pronotal collar tuberculate, punctate. Anterior margin of pronotum often with larger,
yellowish tubercles. Callar region strongly tuberculate, hairs often arising from tubercles.
Anterior half of pronotum tuberculate, tubercles with hairs arising from them; posterior
half strongly punctate. Ear-like lateral expansions extending from pronotum. Posterior
pronotal angles lacking, or only vaguely present. Scutellum deeply punctate. Corium
concolorous with body, membrane darker, venation with little or no anastomosis.
Abdomen wider than pronotum. Connexiva lacking tubercles; pale spots sometimes
present on interior side, margined in dark brown or black on anterior, posterior and lateral
sides. Spur present at posterolateral angle of connexival segments II-VI, sometimes VII;
segment VII with an anterior pale spot. Abdomen lacking ventral markings. Genital
capsule rim straight, entire, margined with hairs. Genital capsule in posterior view
completely punctate, medially rugose. Propleural acetabular suture with strong keel.
Metathoracic scent gland rugose, sometimes with small punctations; anterior auricle often
with pale dorsal spot. Posterior metapleural margin straight. Area surrounding abdominal
segment IV spiracle not raised as trigonal area. Knob present on metacoxa, especially
prevalent in males. Pro- and mesofemora with ventrodistal pair of small spines, followed
by pair of much larger spines, both pairs darker in color than remainder of spines;
26
preceded proximally by 2-3 pairs of spinose tubercles. Metafemora armed dorsally with
spines or spinose tubercles, sometimes in 2-3 lengthwise rows; lateral outer face with
three large flattened tubercles in series; venter with arrangement similar to pro- and
mesofemora, but two distal-most pair of spines fused in each row; more often with spines
than tubercles. Anterior face of metatibiae darker than posterior face, male metatibiae
arcuate; external apical tubercle lacking.


range: 20.0 - 22.7, X = 21.3.

Width at pronotum (mm) – Range: 7.6 - 10.5, X = 9.13; male range: 8.8 - 10.5, X =

9.5; female range: 7.6 - 9.7, X = 8.8.

Width at connexiva (mm) – Range: 10.9 - 14.3, X = 12.3; male range: 10.9 - 14.3, X =

12.6; female range: 11.0 - 12.4, X = 11.9.
Diagnosis – P. auriculatus is unique among Piezogaster, as the only species with a
combination of uniquely shaped pronotal expansions (Fig. 135), dark red to dark reddish-
orange color and the dark margins around its connexival segments.
Distribution – (Fig. 6) Belize (TAMU); El Salvador (NMNH); Guatemala; Honduras
(NMNH); Mexico: Chiapas, Guerrero, Jalisco, Morelos, Nayarit, Oaxaca, Puebla,
Veracruz.
Notes – Brailovsky & Barrera described P. scitus as very similar to P. auriculatus, but
with P. scitus having less pronounced tubercles, a lesser-defined coxal process (I call a
"knob"), and unique-looking parameres. Among the twenty-one specimens I examined,
the first two characters were quite variable, and Brailovsky & Barrera's illustrations of the
27
parameres provide little distinction. Additionally, all specimens from Brailovsky &
Barrera's description of P. scitus originate from locales where P. auriculatus is already
known to exist. Moreover, after examining specimens identified as P. scitus by
Brailovsky and comparing them to specimens identified as P. auriculatus, it appears that
the distinguishing characteristics given for P. scitus by Brailovsky & Barrera (1984) are
not distinct enough to warrant recognition of a separate species. Therefore, I formally
synonymize P. scitus with P. auriculatus.
Moreover, after looking at the historic distributions and comparing them to
distributions of the specimens examined (see Fig. 6), it appears that historic distributions
of P. auriculatus in New Mexico and Texas noted in Uhler (1876) are inaccurate. I was
unable to find any specimens from either New Mexico or Texas, and there is a large
distribution gap separating the remainder of the distribution from these two locations.
Additionally, the localities now known for P. auriculatus are subtropical to tropical,
whereas New Mexico and Texas are more arid regions.
Specimens examined contained one homoeotype from UMC designated and
labeled as such by Thomas Yonke.
Piezogaster basilicus Brailovsky & Barrera

(Fig. 50-53)
Piezogaster basilicus Brailovsky & Barrera 1984; p. 137; Mexico: Veracruz.
Body color cinnamon to dark brown, covered with minute hairs. Head with
tubercles sparse to lacking, postocular area tuberculate, not darker in color than remainder
28
of head. Antennal segment I longest, II, IV subequal, III shortest; ultimate segment
orange to reddish-orange, remainder of segments concolorous with remainder of body.
segments I, II, IV subequal, segment III shortest. Anterior pronotal collar tuberculate,
deeply punctate. Pronotum margined with minute tubercles, callar region impunctate,
remainder heavily punctate; lateral expansions lacking. Posterior pronotal angles lacking.
Scutellum rugosely punctate, yellowish with dark brown margins. Corium concolorous
with body, veins slightly paler; membrane darker, veination with little or no anastomosis.
Pronotum wider than abdomen. Connexiva appearing bicolored, yellowish anteriorly,
cinnamon to dark brown posteriorly; tubercles lacking, hairs minute. Connexival spur on
each segment minute to lacking. Abdomen lacking ventral markings. Genital capsule rim
entire, slightly dorsally depressed, margined with small hairs. Genital capsule in posterior
view medially rugose, entirely punctate. Prothoracic acetabluar suture not keeled.
Auricles of metathoracic scent gland yellowish, sometimes with a darker margin.
Posterior metapleural margin sinuous. In males, raised trigonal area present surrounding
abdominal segment IV spiracle. Metacoxal knob lacking. Pro- and mesofemora generally
unarmed, save the ventrodistal with 2-3 pairs of spines; distal most smallest, penultimate
significantly larger than other spine or spines. Metafemora slightly arcuate; dorsodistal
side with a trigonal arrangemnt of three spinose tubercles, sometimes yellowish in color.
Ventrodistal with minute ultimate spine pair, followed by three pairs of larger spines in
females; males with three spines on outer-face row, the proximal-most spine not equally
spaced with remainder; one spine on inner-face row; ultimate, penultimate pairs
sometimes fused. Anterior face of metatibiae concolorous with posterior face; external
29
apical tubercle present in males. Tarsi orange to reddish-orange in color.
Length (mm) – Male: 24.1; female: 25.4.
Width at pronotum (mm) – Male: 7.2; female: 7.7.
Width at connexiva (mm) – Male: 6.2, female: 7.4.
Diagnosis – P. basilicus is similar in form and color to P. tetricus, with a pronotum wider
or subequal to the abdominal width (Fig. 142), and contrasting bicolored connexival
segments that are easily seen with naked eye. P. basilicus is longer than P. tetricus –
more than 23 mm by my measurements as well as those in Brailovsky & Barrera (1984) –
and is usually proportionately narrower than P. tetricus in the anterior abdominal region as
well.
Distribution – (Fig. 7) Mexico: Chiapas, Colima (TAMU), Morelos, Oaxaca, Veracruz.
Notes – Brailovsky & Barrera (1984) stated that the main difference between P. basilicus
and P. tetricus is that P. tetricus is not longer than 21 mm, whereas P. basilicus is
between 24-25 mm. However, I found specimens of what was apparently P. tetricus that
ranged from 19 - 23.8 mm in length, but did not exceed 24 mm. Moreover, the ranges of
these two species overlap in several places. Lack of an adequate number of identified P.
basilicus specimens prevents me from drawing absolute conclusions, because the main
character separating the two species is a difference in size.

30
Piezogaster bolivianus Brailovsky
Piezogaster bolivianus Brailovsky 1993; p.113; Bolivia.
Like P. acuminatus, P. bolivianus has pronotal angles that extend laterally forming a
spine, as opposed to the anteriorly curving spines of the pronotal angles of P. dilatatus.
However, like P. dilatatus, P. bolivianus has the ultimate antennal segment unicolored,
not bicolored like P. acuminatus. Moreover, the pronotal angles of P. acuminatus are
described as being thinner, and the tubercles present on the femora and tibia are not as
dense or robust as those of P. bolivianus. I have not seen an actual specimen of this
species, but based on the descriptions and figures in Brailovsky (1993), compared to the
specimens I have examined, it appears that this species is still valid.
Measurements – From Brailovsky (1993). Males: n = 1; females: n = 1.
Distribution – (Fig. 8) Bolivia.
Piezogaster calcarator (Fabricius)

(Fig. 54-57)
Coreus calcarator Fabricius 1803; p. 192; United States: "Carolina".

Archimerus calcarator: Stål 1870; p.137.
Piezogaster calcarator: O'Shea 1980; p. 214. In part.
Piezogaster calcarator: Baranowski & Slater 1986; p. 39.
Piezogaster calcarator: Henry & Froeschner 1988; p. 88. In part.
Body color pale brown, often with an orange hue; covered with minute white hairs.
31
Head tuberculate, remainder of body tubercles sparse. Postocular area sometimes darker
than remainder of head, always tuberculate. Antenna with erect hairs, sometimes dark
brown or black; segments unicolorous. Antennal segment I, tuberculate, longest; segment
II, IV subequal, segment III shortest; all segments concolorous. Laterally projecting jugal
shelf present beneath base of antennal segment I. Beak segments I, II, IV subequal in
length, segment III shortest. Pronotal collar punctate, deeply so laterally. Anterior
margin of pronotum tuberculate; callar region often with hairs, impunctate; remainder of
pronotum deeply punctate. Pronotal lateral expansions lacking. Posterior pronotal angles
present. Scutellum rugosely punctate. Corium usually concolorous with body, punctate;
membrane apparently somewhat darker; venation slightly anastomizing. Abdomen wider
than pronotum. Connexival segments unicolorous with minute hairs, tuberculate;
posterolateral spur on each segment faint to lacking, if present, at segments IV, V, VI.
Abdomen lacking ventral markings. Rim of genital capsule entire, but slightly depressed
dorsally. Seen from posterior view, genital capsule medially rugose, punctate. Propleural
acetabular suture lacking keel. Posterior metapleural margin sinuous. Area surrounding
abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob lacking. Pro-
and mesofemora each with pair of ultimate minute ventrodistal spines preceded proximally
by much larger penultimate pair, preceded by 3-4 pairs of tubercles. Dorsal area of
metafemora sometimes covered with minute tubercles. Venter of femora with 5-6 pairs of
spines or spinose tubercles; distal-most pair each with doubled point. Spine on interior
row of ventro-metafemora usually significantly larger than remainder of spines. Anterior
face of metatibiae concolorous with posterior face, male metatibiae arcuate; external
apical tubercle lacking.

32

Length (mm) – Range: 17.7 - 21.0, X=19.5; male range: 17.9 - 21.0, X=19.9; female

range: 17.7 - 20.9, X = 19.1.


female range: 6.3 - 7.0, X = 6.7.


female range: 8.4 - 9.6, X = 9.0.
Diagnosis – Morphologically, P. calcarator is similar to both P. alternatus and P.
indecorus, however, the body color is much paler than the latter two, often with an
orange hue and all of the connexival segments of P. calcarator are unicolorous, as
opposed to the bicolored segments of P. alternatus and P. indecorus. P. calcarator does
not have the dark brown to black hairs covering its body that P. alternatus does.
Additionally, P. calcarator occupies only the southeast region of the United States,
whereas P. alternatus and P. indecorus are more cosmopolitan across the United States
and Mexico, respectively.
Distribution – (Fig. 9) United States: Alabama (CU), Florida, Georgia (NMNH),
Mississippi (CAS) North Carolina, South Carolina (Fabricius (1803) refers to
"Carolina").
Notes – See P. alternatus notes regarging the reinstatement of P. alternatus.
One homoeotype examined from UMC designated and labeled as such by Thomas
Yonke.
33
Piezogaster camposi (Montandon)

(Fig. 58-61)
Archimerus camposi Montandon 1897; p. 246; Ecuador.

Sephina humeralis Distant 1901; p.420; New synonymy.
Piezogaster camposi: O'Shea 1980; p. 214.
Piezogaster humeralis: O'Shea 1980; p. 214.
Body color reddish-orange to orange with brownish-black to black accents and
extremities. Head hairless, lacking tubercles; remainder of body with minute hairs,
tubercles lacking. Antenniferous tubercles with anterior margin darker than remainder of
head. Postocular area not darker than remainder of head or tuberculate. Each antennal
segment shorter than previous segment as series extends distally; all segments
concolorous. Laterally projecting jugal shelf lacking beneath base of antennal segment I.
Beak segments I, II, IV all subequal, segment III shortest. Anterior margin of pronotum
tuberculate up to lateral expansions; margins of expansions not tuberculate. Anterior
pronotal collar punctate, laterally tuberculate. Anterior surface of pronotum dark brown
to black, callar region reddish-orange to orange, impunctate, hairless and lacking
tubercles; posterior half strongly punctate with three dark brown to black longitudinal
stripes; posterior margin dark brown to black. Pronotum with rounded lateral expansions.
Posterior pronotal angles present, but quite obtuse. Scutellum rugosely punctate, darkly
margined. Each corium with dark brown to black macula; membrane slightly lighter in
color than remainder of body, venation with little or no anastomosis. Pronotum wider
than abdomen, abdomen sometimes wider in females. Connexival segments reddish-
orange anteriorly, brownish-black to black posteriorly, each segment usually with more
reddish-orange than brownish-black to black. Posterolateral spur present on connexival

34
segments III-VI, sometimes II, spurs enlarged in males. Abdomen patterned with dark
brown to black on orange. Genital capsule rim, bilobate from dorsal view, not dorsally
depressed, margined with minute hairs. Surface of genital capsule in posterior view
medially rugosely punctate. Propleural acetabular suture lacking keel. Each thoracic
pleuron dark brown to black with an orangish-red spot. Area surrounding metathoracic
scent gland rugose, pale orange. Posterior metapleural margin straight. Area surrounding
and mesofemora with ventrodistal pair of minute ultimate spines; penultimate spines much
larger. Male metafemora arcuate, having sparsely distributed spinose tubercles. Venter of
metafemora with 4-5 pairs of spinose tubercles running the length of the metafemora,
medial inner spinose tubercle usually largest; ventrodistal spines minute. Male metatibiae
concolorous, arcuate; external apical tubercle lacking.

Length (mm) – Range: 19.4 - 21.2, X = 20.5; male: 20.2; female range: 19.4 - 21.2, X =
20.6.

Width at pronotum (mm) – Range: 8.0 - 8.9, X = 8.4; male: 8.2; female range: 8.0 - 8.9,
X = 8.5.

Width at connexiva (mm) – Range: 7.4 - 9.2, X = 8.3; male: 7.4; female range: 8.0 - 9.2,
X = 8.5.
Diagnosis – This species is easily recognized and differentiated form other Piezogaster by
the unique reddish-orange and brownish-black to black striped pattern on the pronotum
and the shape of the pronotal lateral expansions (Fig. 58, 60). Also, the dark brown to
black macula on each hemelytron makes it easily discernible from other Piezogaster
35
species (Fig. 58, 60).
Distribution – (Fig. 10) Ecuador, Venezuela (BMNH).
Notes – O'Shea (1974) placed Sephina humeralis Distant in the genus Archimerus and
suggested that it would prove to be a synonym of (then) Archimerus camposi, however,
O'Shea did not deal with this pending synonymy in his subsequent work (1980). I had the
opportunity to examine a specimen labeled Sephina humeralis from Distant's collection at
the British Museum and have found it to be identical to P. camposi, having the unique
reddish-orange and brownish-black to black striped pattern on the pronotum, uniquely
expanded pronotum, and the dark brown to black macula on the leathery portion of each
hemelytron. The original descriptions of both species mention all of these characters and
the body lengths and pronotal widths of both types are within 2 mm of each other.
Apparently, O'Shea indeed was correct. Therefore, I formally synonymize P. humeralis
with P. camposi, the elder name of the two.
Piezogaster chiriquinus (Distant)

(Fig. 62-65)
Archimerus chiriquinus Distant 1893; p. 355; Panama.

Archimerus chiriquiensis: Lethierry & Severin 1894; p17.
Piezogaster chiriquinus: O'Shea 1980; p. 214.
Body color cinnamon to ruddy brown, covered with minute hairs. Postocular area of
head darker than remainder of head, tuberculate. Antennal segment I longest, segments
II, IV subequal, segment III shortest; ultimate segment usually reddish-orange, other
segments concolorus with body. Laterally projecting jugal shelf slightly visible beneath
36
base of antennal segment I. Beak segments I, II, IV subequal, segment III shortest.
Pronotal collar rugosely punctate, especially on lateral sides. Anterior half of pronotum
with minute tubercles. Callar region with two distinct impunctate regions, usually
tuberculate with some hairs. Rest of pronotal surface heavily punctate. Pronotum slightly
expanded laterally. Posterior pronotal angles present. Scutellum usually paler in color,
often yellowish, heavily punctate, slightly rugose, usually darkly margined. Corium
concolorous with body, membrane darker than remainder of body; veins of membrane
anastomosing. Abdomen subequal in width to pronotum. Connexiva lacking tubercles,
each usually with a pale anterior patch. Spur present at posterolateral angle of connexival
segments IV, V, VI in males, females sometimes lacking spurs. Pair of small, pale spots
present ventrally on abdominal segments III-VI, sometimes VII. Genital capsule rim
sometimes curving gently ventrally, entire, margined with hairs; rim extends posteriorly as
to form a shelf or lip. In posterior view, genital capsule medially rugose; covered entirely
with minute punctations. Propleural acetabular suture lacking keel. Posterior metapleural
margin sinuous. Area surrounding abdominal segment IV spiracle surrounded by raised
trigonal area. Metacoxal knob lacking. Ventrodistal end of pro- and mesofemora with
two darker spines; distal spine much smaller, preceded proximally by 3-4 pairs of
tubercles, sometimes pale. Venter of metafemora with two longitudinal rows of spinose
tubercles, ending distally with a darker trigonal spine, often with double-point. Dorsal of
metafemora with 3-4 very flattened tubercles similar in color to lighter regions of
metafemora; anterior spotted with tubercles; in males, three medially located spines
present, middle spine significantly smaller. Anterior face of metatibiae often darker than
posterior face. In males, metatibiae noticeably arcuate; external apical tubercle present.
37
All tarsi similar in color to antennal segment IV.

Length (mm) – Range: 18.6 - 21.7, X=19.9; male range: 18.6 - 21.7, X = 20.4; female

range: 18.7 - 21.1, X = 19.5.


female range: 7.0 - 7.9, X = 7.4.


female range: 6.4 - 8.1, X = 7.1.
Diagnosis – P. chiriquinus stands out from similar species by its short, rounded, lateral
expansions on its uniquely shaped pronotum (Fig. 134) and the reddish-orange ultimate
tarsal and antennal segment. In males, P. chiriquinus can be further discerned by the
arrangement of two large spines separated by a much smaller spine on the dorsal side of
the metafemora.
P. chiriquinus is similar in shape to P. odiosus, P. tetricus, appearing elongate,
with the pronotum being wider than the abdomen (Fig. 142). P. chiriquinus is more
tuberculate than both P. odiosus and P. tetricus. Compared to P. scutellaris, P.
chiriquinus is longer and a bit wider, with less incrassate metafemora than P. scutellaris,
and also lacks the pale yellow colored metafemoral tubercles found on P. scutellaris. P.
odiosus, P. tetricus, and P. scutellaris all lack the unique pronotal expansions of P.
chiriquinus (Fig. 134).
Distribution – (Fig. 11) Costa Rica (NMNH); Honduras (UCB); Mexico: Chiapas
(TAMU), Jalisco (NMNH), Morelos, Oaxaca (TAMU); Panama.

38
Piezogaster chontalensis (Distant)

(Fig. 66-69)
Capaneus chontalensis Distant 1893; p. 354; Nicaragua.

Piezogaster chontalensis: O'Shea 1980; p. 214.
Body color brown to orangish-brown, femora and pronotum a darker chestnut color;
covered with minute hairs, appearing velvety on pleura and sterna of thorax. Body
lacking prominent tubercles. Postocular area concolorous with head, tubercles lacking.
Antennal segments, I, II subequal, longest, segment III shorter, segment IV shortest; all
segments concolorous. Segment II, III with hairs often in parallel rows running
lengthwise. Laterally projecting jugal shelf present beneath base of antennal segment I.
Beak segments I, II, III subequal, IV longest. Pronotal collar tomentose, lacking
tubercles. Anterior pronotal margin lacking tubercles, surface with minute punctations.
Callar region two distinct impunctate areas, lacking tubercles, often with patches of
tomentose hairs. Lateral pronotal expansions acuminating in a point, slightly turned
anteriorly. Posterior pronotal angles lacking. Scutellum rugosely punctate with corners
of angles often a dirty yellow, glaberous. Corium concolorous with body, membrane
sometimes slightly paler in color; venation with little or no anastomosis. Pronotum wider
than abdomen. Connexiva lacking tubercles, unicolorous; spur present on segments IV,
V, VI. Abdomen lacking ventral markings. Genital capsule rim entire, slightly dorsally
depressed, minute hair on rim sometimes lacking. Surface of genital capsule in posterior
view medially rugose, punctate, depressed between raised area and rim. Propleural
acetabular suture with slight keel. Posterior metapleural margin slightly sinuous. Area
39
lacking. Pro- and mesofemora lacking tubercles, spines. Protibiae, mesotibiae only
vaguely triquetrous in cross-section, nearly round. External groove running length of
protibiae, mesotibiae only vaguely present. Venter of male metafemora with two
lengthwise rows of spinose tubercles, 5-7 to a row, inner medial spinose tubercle usually
largest. Metatibiae concolorous, male metatibiae not arcuate. Distal end of tibiae lacking
fringe of hairs; external apical tubercle lacking.

Length (mm) – Range: 15.3 - 18.9, X = 17.3; males 15.3 and 17.5; females: 17.6 and
19.9.

Width at pronotum (mm) – Range: 5.3 - 7.7, X = 6.6; males 6.4 and 7.0; females: 5.3
and 7.7.

Width at connexiva (mm) – Range: 4.9 - 6.5, X = 6.0; males 4.9 and 6.4; females: 6.0
and 6.5.
Diagnosis – The specimens of P. chontalensis that I examined have pronotal expansions
directed slightly posteriorly at the apex (Fig.129). Comparatively, the expansions of P.
reclusus appear similar, but are turned slightly forward (Fig. 127). P. achillelus has
pronotal expansions that are longer and much more forward swept (Fig. 126) and P.
achilles specimens have expansions that extend laterally as much as they do forward (Fig.
128). P. chontalensis is separated from P. achilles and P. reclusus by the darker chestnut
coloration on the femora and pronotum, which both of the latter lack.
Distribution – (Fig. 12) Nicaragua; Panama (BMNH).
Notes – Two of the specimens examined were from Distant's collection at the British
Museum, one of which was identified by Distant in 1911. Label data show one specimen
found on Acacia sp.

40
Piezogaster congruus Brailovsky & Barrera
Piezogaster congruus Brailovsky & Barrera 1983; p. 71; Peru.
Brailovsky & Barrera stated that P. congruus appears similar in shape and color to P.
rubropictus, and their figures confirm this. Both are brownish-black to black in color and
lack pronotal expansions. However, instead of the red to reddish-orange pronotal stripes
and bicolored connexiva of P. rubropictus, P. congruus has a single ocherous-orange
medial longitudinal stripe extending from the head through the scutellum, which is unique
among Piezogaster species, and connexiva that are completely orange. Brailovsky &
Barrera also noted the presence of an unmistakable finger or tongue-like projection arising
from end of the male's genital capsule (Fig.141), which is unique to the genus. Although I
have not seen a specimen of this species, the description from Brailovsky & Barrera
(1983) points out enough unique characters to easily separate this species from the
remainder of Piezogaster.
Measurments – From Brailovsky & Barrera (1983). Males: n = 1; females: n = 1.
Diagnosis – The unique ocherous-orange medial longitudinal stripe extending from the
head through the scutellum separates P. congruus from all other Piezogaster species.
Distribution – (Fig. 13) Peru.

41
Piezogaster dilatatus (Dallas)

(Fig. 70-73)
Archimerus dilatatus Dallas 1852; p. 418; Venezuela.

Piezogaster dilatatus: O'Shea 1980; p. 214.
Body color pale to medium brown, covered with minute hairs, tubercles. Head
strongly tuberculate. Postocular area darker than remainder of head, tuberculate.
Antennal segment IV longest, segment I, II subequal, segment III shortest; all segments
concolorous. Lacking laterally projecting jugal shelf beneath base of antennal segment I.
Beak segments I, II, IV subequal in length, segment III shortest. Collar of pronotum
punctate, tuberculate. Anterior pronotal margin with dentate tubercles. Callar region
strongly tuberculate, remainder of pronotum strongly punctate, scattered with tubercles.
Pronotum expanded with an anterolateral spine. Posterior pronotal angles present.
Scutellum rugosely punctate, darkly margined. Hemelytra concolorous with body,
membrane with strongly anastomosing venation. Abdomen wider than pronotum.
Connexiva with small hairs, tubercles; ech segment with small, lateral, yellowish patch
present just anterior to posterolateral spur, spur present on connexival segments III-VI,
sometimes II. Venter of abdomen tuberculate; slight raised discoid areas present on
segments IV, V, and sometimes III in males, but present only on segment IV in females.
Genital capsule rim with two dorsal teeth, rim margined with hairs. Genital capsule in
posterior view with minutely granular surface, darkly mottled. Propleural acetabular
suture not keeled. Thoracic pleura punctate, tuberculate. Metathoracic scent gland
auricles yellowish in color. Posterior metapleural margin sinuous. Area surrounding
42
and mesofemora covered dorsally, laterally with tubercles, sometimes in lengthwise rows;
ventrally with 4-5 pairs of spines or spinose tubercles, getting smaller as series runs
proximally. Dorsal, lateral of metafemora with spines or spinose tubercles sometimes in
series; ventrally with 4-5 pairs of spines or spinose tubercles, with distal most pairs often
fused together. Anterior face of metatibiae concolorous with posterior face; metatibiae
not arcuate, lacking external apical tubercle.


range: 18.6 - 20.5, X = 21.4.


female range: 7.0 - 8.0, X = 7.5.


female range: 7.3 - 11.2, X = 10.5.
Diagnosis – This species is similar to P. obscuratus, both of which have an abdomen
wider than the pronotum, but P. dilatatus lacks bicolored connexiva and is covered with
many more tubercles. P. dilatatus also has a prominent anterolateral spine on its pronotal
expansions (Fig. 70, 72, 80, 82), and lacks the pale yellow pleural fascia of P. obscuratus
(Fig. 71, 73, 81, 83). In addition, P. dilatatus possesses large, yellowish, discoid areas on
the venter of its abdomen, which P. obscuratus lacks, and the genital capsule of P.
dilatatus is toothed and has a rough, mottled appearance, whereas the genital capsule of
P. obscuratus is entire and is medially rugose and punctate.
Distribution – (Fig. 14) Bolivia; Brazil; Ecuador; Peru; Venezuela.

43
Piezogaster humerosus (Distant)

(Fig. 74-75)
Capaneus humerosus Distant, 1893; p. 354; Mexico.

Piezogaster humerosus: O'Shea 1980; p. 214.
I received only one badly damaged female specimen identified as P. humerosus,
which was not enough material to write a complete description, however, the material I
had matched the original description by Distant (1893), and subsequent description by
Brailovsky and Barrera (1984). In the literature, P. humerosus is described as having
broadly produced, somewhat pointed lateral expansions of the pronotum (Fig. 131), with
black pronotal tubercles that mainly occur on the expansions. I found these tubercles to
be strong, rounded, reddish-brown to black in form and color. P. humerosus also has a
broken, reddish-brown fascia on its pleura (Fig. 75), which I was able to identify as well.
Brailovsky & Barrera (1984) noted that this species is rarely seen or collected in its
endemic range in Tamaulipas and Veracruz, Mexico.
Measurements – Females: n = 1.
Length (mm) – 25.5.
Width at pronotum (mm) – 13.1.
Width at abdomen (mm) – 12.0.
Distribution – (Fig. 15) Mexico: Tamaulipas, Veracruz.

44
Piezogaster indecorus (Walker)

(Fig. 76-79)
Archimerus indecorus Walker, 1871; p. 64. Mexico.

Archimerus herrichi Blöte 1938. New synonymy.
Piezogaster herrichi: O'Shea 1980; p. 214.
Piezogaster indecorus: O'Shea 1980; p. 214.
Body color variable; pale light brown to very dark brown. Minute hairs covering
most of body. Head tuberculate, especially on tylus. Postocular area often darker than
remainder of head, tuberculate. Antennal segment I longest; segment II, IV subequal;
segment III shortest; segments I, II, III with short dark brown to black hairs, ultimate
segment sometimes lighter in color. Laterally projecting jugal shelf present beneath base
of antennal segment I, but minute. Beak segment I longest, segments II, IV subequal,
segment III shortest. Anterior pronotal collar strongly punctate, sometimes tuberculate.
Anterior pronotal margin tuberculate. Callar region with hairs, impunctate; remainder of
pronotal surface strongly tuberculate; lateral expansions lacking. Posterior pronotal
angles present. Scutellum sometimes yellowish, somewhat rugose, deeply punctate.
Corium concolorous with body, membrane usually darker, venation slightly anastomosing.
Pronotum subequal in width to abdomen. Connexival segments with minute hairs,
tuberculate; each segment with a pale yellow spot anteriorly, spot lacking dark margin
laterally, but connexiva darkly margined on lateral margins posterior to the spot.
Connexival posterolateral spurs minute to lacking. Pair of minute, yellowish,
ventroabdominal spots often present on segments III-VI. Genital capsule rim entire but
dorsally depressed; margined dorsally with minute hairs. Genital capsule in posterior view
medially rugose, punctate. Propleural acetabular suture with strong keel. Thoracic pleura
45
punctate, sometimes with tubercles. Posterior metapleural margin sinuous. Area
lacking. Pro- and mesofemora with ventrodistal pair of minute ultimate spines;
penultimate spines much larger, often preceded by 3-4 pairs of pale tubercles or minute
spines. Dorsum of metafemora with spinose tubercles, often in lengthwise rows. Venter
of metafemora with similar arrangement to pro- and mesofemora except two most distal
pairs of spines fused; preceded proximally by spines more often than tubercles. In males,
single ventral spine on posterior of metafemora significantly larger than remainder of
metafemoral spines. Interior face of metatibiae often darker than exterior face; in males,
metatibiae arcuate; external apical tubercle lacking.


range: 17.6 - 21.6, X = 19.3.


female range: 6.3 - 8.0, X = 7.2.


female range: 6.7 - 10.0, X = 7.9.
Diagnosis – This extremely variable species has been confused with P. alternatus in
collections and in areas where the two species are sympatric, sharing almost every visible
character except the white and scattered black erect hairs on the head, pronotum, and
femora, which P. indecorus lacks. Additionally, antennal segments II and III of P.
indecorus often do not appear as robust as those of P. alternatus because of the lack of
white and scattered black erect hairs. P. indecorus is also a southern species, limited to
46
Mexico and the southwestern United States, whereas P. alternatus is cosmopolitan over
most of the United States east of the Continental Divide.
Distribution – (Fig. 16) El Salvador (NMNH); Guatemala (NMNH); Honduras
(TAMU); Mexico: Chiapas (TAMU), Chihuahua (UMC), Coahulia (UCB), Distrito
Federal, Durango, Estado De Mexico, Guerrero, Hidalgo, Jalisco (UCB), Michoacan,
Morelos, Nuevo Leon (TAMU), Oaxaca, Puebla, Sinaloa (UCB), Tlaxcala, Veracruz;
United States: Arizona, New Mexico (TAMU), Texas (TAMU).
Notes – Many specimens that I initially identified as P. indecorus, I later found to have
tags identifying them as P. herrichi. As a result, I examined the original description of
Archimerus squallus Herrich-Schäffer (1842), the designated name-bearer for P. herrichi
as described in Blöte (1938), and determined that the original description is quite vague
and provides no characteristics for distinguishing P. herrichi from P. indecorus.
Brailovsky & Barrera (1984) stated that the species differs from P. indecorus in that
connexiva IV and V do not possess the pale yellow anterior spot that P. indecorus does.
However, after examining hundreds of specimens assigned to both taxa, I deemed this
differentiation too variable. Moreover, the smaller range of P. herrichi is within the range
of P. indecorus (Fig. 30). Taking all of this into consideration, I formally synonymize P.
herrichi with P. indecorus.
Piezogaster loricata (Distant)

(Fig. 122)
Ojedana loricata Distant 1893; p. 356; Panama. Monotypic.

Piezogaster loricata: O'Shea 1980; p. 214.
47
The only available records of this species exist in Distant's original publication, which
was noted and carried over into O'Shea (1980) when he synonymized the genus Ojedana
with Piezogaster. Moreover, P. loricata is the only species of this now defunct genus.
Only one known specimen was cited in the in Distant's type description, and I was not able
to obtain any specimens for examination.
Distant (1893) described P. loricata as black with a broad, oblique sternal fascia and
eight pale yellow abdominal discoid areas running in paired series. According to Distant's
illustration and description, P. loricata also bears subacute pronotal angles similar to P.
obscuratus (Fig. 133). This species might be also be similar to P. dilatatus, which has
abdominal discoid areas as well.
Measurements – From Distant (1893). n = 1
Length (mm) – 21
Width at pronotum (mm) – 10
Distribution – (Fig. 17) Panama.
Piezogaster multispinus (Stål)

(Fig. 123)
Capaneus multispinus Stål 1862; p. 280; Mexico.

Piezogaster multispinus: O'Shea 1980; p. 214.
I have not had the opportunity to eaxamine a specimen of P. multispinus, and the
descriptions I have seen for this species lack detail. However, the photos and illustrations
depict an anteriorly directed and expanded pronotum with an extremely dentate-serrate
margin, as well as multispinose femora. The dentate-serrate pronotal margin (Fig. 123)
48
alone is unique for this genus, and is sufficient to separate P. multipinus from the
remainder of Piezogaster.
Measurements – From Stål (1862). n = 1
Length (mm) – 20
Width (mm) – 6
Distribution – (Fig. 18) Mexico: Oaxaca, Veracruz.
Piezogaster obscuratus (Montandon)

(Fig. 80-83)
Capaneus obscuratus Montandon 1899; p. 191; Ecuador.

Piezogaster obscuratus: O'Shea 1980; p. 214.
Body color pale to dark brown, covered with minute hairs. On head, postocular area
seldom darker than remainder of head, or tuberculate. Antennal segment I longest,
segment IV shorter than segment I, segment II shorter than segment IV, segment III
shortest; all segments concolorous. Laterally projecting jugal shelf present beneath base
of antennal segment I. Beak segments I, II, IV subequal, segment III shortest. Anterior
pronotal collar punctate. Anterior pronotal margins with sparse dentate tubercles; callar
region impunctate with minute, sometimes decumbent hairs, remainder of pronotum
strongly punctate. Pronotum expanded laterally, lateral angles subacute. Posterior
pronotal angles lacking. Surface of scutellar angles smooth, usually yellowish in color,
remainder of scutellum rugosely punctate. Hemelytra concolorous with body, membrane
with some veins anastomosing. Abdomen wider than pronotum. Connexival segments
49
with minute hairs, tubercles lacking; anterolateral angles of each segment yellowish in
color; posterolateral spur present on segments IV-VI, sometimes III. Ventral abdominal
segments III-VI each with a pair of small, pale, usually yellow spots. Genital capsule rim
entire, but slightly depressed dorsally, margined with hairs. In posterior view, genital
capsule medially rugose, punctate. Smooth yellowish fascia present on on each thoracic
pleura. Propleural acetabular suture not keeled. Posterior metapleural margin straight.
Area surrounding abdominal segment IV spiracle not raised as trigonal area. Metacoxae
with prominent lateral hooked knob. Pro- and mesofemora with pair of ventrodistal
spines often preceded by 2-3 pairs of smaller spines. Metafemora covered dorsally and
laterally with small sometimes spinose tubercles, these sometimes in lengthwise rows.
Metafemora with pair of double-pointed spines ventrodistally, preceded by 2-3 pairs of
spinose tubercles. Anterior face of metatibiae concolorous with posterior face. In males,
metatibiae somewhat arcuate; external apical tubercle lacking.


range: 18.6 - 22.5, X = 20.6.


female range: 7.7 - 10.3, X = 9.1.


female range: 8.6 - 10.6, X = 10.0.
Diagnosis – P. obscuratus is similar in appearance to P. dilatatus, both having an
abdomen wider than the pronotum. However, P. obscuratus has bicolored connexiva, a
yellowish fascia on each of the thoracic pleura (Fig. 81, 83), and lacks the anterolateral
50
spine on the pronotal expansions, which P. dilatatus has (Fig. 70, 72, 80, 82). P.
obscuratus males also have an entire genital capsule, unlike the toothed genital capsule of
P. dilatatus.
Distribution – (Fig. 19) Ecuador.
Notes – Two specimens intercepted in Pennsylvania on bananas were examined. An
additional specimen was found on Algarrobo sp.
Piezogaster odiosus (Stål)

(Fig. 84-89)
Capaneus odiosus Stål, 1862; p. 291; Mexico.

Capaneus dolosus Walker 1871. Synonymized by Distant (1881).
Piezogaster odiosus: O'Shea 1980; p. 214.
Body color brown to dark brown, or nearly black, covered with minute hairs.
Head sparsely tuberculate to lacking. Postocular area as dark as remainder of head, with
paler tubercles. Antennal segment I longest. Segments, II, IV subequal, segment III
shortest; ultimate antennal segment usually reddish-orange in color. Laterally projecting
jugal shelf present beneath base of antennal segment I. Beak segments I, II, IV subequal,
segment III shortest. Anterior pronotal collar strongly punctate. Anterior margins of
pronotum with minute tubercles, callar region often tuberculate, impunctate, remainder of
pronotum punctate; lateral expansions lacking. Posterior pronotal angles lacking.
Scutellum rugosely punctate. Corium concolorous with body, membrane sometimes
apparently darker than remainder of the body, venation with little or no anastomosis.
Pronotum wider than abdomen in males, subequal in females. Connexiva with minute
51
hairs, unicolorous, tubercles lacking. Minute posterolateral spur present on connexival
segments IV-VI, sometimes also on III. Abdomen lacking ventral markings. Genital
capsule rim entire, sometimes depressed dorsally. Seen from posterior view, genital
capsule medially rugose, punctate. Propleural acetabular suture not keeled. Posterior
metapleural margin straight. In males, abdominal segment IV spiracle surrounded by a
raised ventrally-pointing trigonal area. Metacoxal knob lacking. Venter of pro- and
mesofemora with ventrodistal pair of minute ultimate spines; penultimate spines much
larger, preceded proximally by 2-3 pairs of sometimes spinose tubercles. Dorsal and
lateral surfaces of metafemora marked sparsely with spinose tubercles. Male metafemora
slightly arcuate, with same spinose pattern ventrally as pro- and mesofemora, but most
distal spines often fused into single double-pointed spine; females with up to five pairs of
spines or spinose tubercles proximally following double-pointed spine. Anterior face of
metatibiae concolorous with posterior face; male metatibiae arcuate with external apical
tubercle. All tarsi usually same reddish-orange color as antennal segment IV.


range: 16.7 - 22.8, X = 19.9.


female range: 5.3 - 7.3, X = 6.6.


female range: 4.1 - 7.5, X = 6.4.
Diagnosis – P. odiosus is very similar morphologically to P. tetricus, P. rubronotatus, and
P. vates, all of which lack pronotal expansions, and have a pronotum wider or subequal to
52
the width of the abdomen (Fig. 142). However, P. odiosus usually is darker in color than
P. tetricus and lacks the pale anterolateral spot on each connexival segment. P. odiosus
also has the reddish-orange tarsi which P. tetricus lacks. P. vates appears nearly identical
to P. odiosus, but the body of P. vates is longer (more than 23 mm, whereas P. odiosus is
shorter than 23 mm, according to my measurements and those of Brailovsky & Barrera
(1984)). P. odiosus also lacks the red semilunar pronotal band of P. rubronotatus, and is
shorter in length (less than 23 mm).
Distribution – (Fig. 20) Belize (TAMU); Costa Rica (UMC); Guatemala; Honduras;
Jamaica (NMNH); Mexico: Chiapas, Colima (NMNH), Guerrero, Jalisco, Morelos,
Oaxaca, Puebla (UMC), Quintana Roo, San Luis Potosi, Tabasco, Tamaulipas, Veracruz,
Yucatan; Nicaragua; Panama; Venezuela (NMNH).
Notes – I examined two homoeotypes from UMC designated and labeled as such by
Thomas Yonke. One specimen examined was intercepted in Louisiana on a bunch of
bananas. Additionally, Schaefer & O'Shea (1979) noted Citrus sinensis as a host plant
and Schaefer & Mitchell (1983) noted Pachyrrizus sp. as a host plant.
Piezogaster reclusus Brailovsky & Barrera

(Fig. 90-93)
Piezogaster reclusus Brailovsky & Barrera, 2000; p. 278; Costa Rica.
Body color yellowish to orangish-brown, covered with minute, almost velvety, hairs
on pleura, venter of thorax, tubercles sparse to lacking. Postocular area never darker than
remainder of head, slightly tuberculate. Antennal segment I longest, II, IV subequal, III
53
shortest; all segments concolorous; segment II, III with hairs in parallel rows running
lengthwise. Lacking laterally projecting jugal shelf beneath base of antennal segment I.
Beak segment IV longest, I, II subequal, segment III shortest. Pronotal collar impunctate,
rarely tuberculate. Pronotum margined with sparse tubercles, surface heavily punctate.
Callar region lacking tubercles or punctation. Pronotum with prominent lateral
expansions pointing anterolaterally. Posterior pronotal angles lacking. Scutellum
punctate except at anterolateral angles. Corium concolorous with body; membrane
darker, venation rarely anastomosing. Pronotum wider than abdomen in males; variable in
females. Connexiva lacking tubercles, unicolorous. Minute posterolateral connexival spur
present on segments VI, VII, sometimes V. Abdomen lacking ventral markings. Genital
capsule rim straight, entire, with sparse hairs. Genital capsule in posterior view rugose,
covered with punctations, sometimes darker than remainder of the genital capsule.
Propleural acetabular suture not keeled. Posterior metapleural margin straight. Area
lacking. Profemora, mesofemora with 1-2 pairs of slight to vague tubercles on distal
ventral side. Protibiae, mesotibiae vaguely triquetrous in cross-section, nearly round.
External groove running length of protibiae and mesotibiae only vaguely present. Distal
end of tibiae lacking fringe of hairs. Three to four pairs of darkened spines running
ventral length of metafemora, becoming less acute distally; spine pairs more prominent in
males. Male metatibiae arcuate, anterior face of metatibiae concolorous with posterior
face; external apical tubercle lacking.

54


range: 16.2 - 18.9, X = 17.7.


female range: 5.8 - 7.9, X = 6.9.


female range: 6.0 – 6.8, X = 6.3.
Diagnosis – When compared to P. achillelus, P. achilles and P. chontalensis, P. reclusus
has pointed pronotal expansions that are turned slightly forward (Fig. 127). The
expansions of P. chontalensis appear most similar to P. reclusus, however they are
directed slightly posteriorly at the apex (Fig. 129). The pronotal expansions of P.
achillelus are longer and much more forward swept (Fig. 126), while P. achilles has
expansions that extend laterally as much as they do forward (Fig. 128).
Distribution – (Fig. 21) Costa Rica.
Notes – Brailovsky & Barrera (2000) stated that this species is similar to both P.
achillelus and P. chontalensis, but is distinguished by its "dark yellow ambarine
coloration," and bright orange to orangish-yellow connexiva, however the connexiva of
the specimens I examined are very similar in color to the remainder of the body overall.
Brailovsky & Barrera also made reference to an "upward and outward" shape to the
pronotal angles, which I interpret as being turned slightly forward (Fig. 127).
Collection data includes specimens found on Acacia costaricensis and A. collinsii.

55
Piezogaster rubronotatus (Stål)

(Fig. 94-97)
Capaneus rubronotatus Stål, 1862; p. 290; Mexico.

Piezogaster rubronotatus: O'Shea 1980; p. 214.
Body color black, occasionally tinged with red, all markings red, body covered with
minute white hairs. Head with slightly tuberculate postocular area. Antennal segment I
longest, segments II, IV subequal, III shortest; all segments concolorous. Lacking
laterally projecting jugal shelf beneath base of antennal segment I. Beak segments I, II, IV
subequal, segment III shortest. Anterior pronotal collar rugosely punctate. Anterior
margin of pronotum with small tubercles, surface heavily punctate. Pronotum with red
semilunar band anteriorly, extending to lateral angles of pronotum. Callar region sparsely
punctate, sparsely covered with small tubercles, hairs. Pronotal expansions lacking.
Posterior pronotal angles lacking. Scutellum punctate, dark brown margins on posterior
sides. Hemelytra concolorous with body, membrane venation with little or no
anastomosis. Pronotum subequal in width to abdomen. Connexiva lacking tubercles,
anterior third to half red to reddish-orange; spur present at posterolateral angles of
connexival segments III-VI, sometimes II. Pair of red to reddish-orange patches on each
visible abdominal segment through VII; patches on VII nearly fused. Genital capsule rim
straight, entire, margined with hairs. Genital capsule in posterior view rugose from
median, turning smooth as progressing laterally. Propleural acetabular suture lacking keel.
Red to reddish-orange band bisecting pro- and mesothoracic pleura, extending diagonally
to posterodorsal corner of metathoracic pleura. Posterior metapleural margin straight. In
males, area surrounding abdominal segment spiracle IV raised, trigonal, pointing ventrally.
56
No metacoxal knob present. Venter of pro- and mesofemora with ventrodistal pair of
minute ultimate spines; penultimate spines much larger, preceded proximally by smaller
spine. Penultimate spine sometimes fused with smaller proximate spine. Dorsodistal side
of metafemora with three small spines arranged trigonally; ventrodistal with pair of large
trigonal spines followed by 2-3 pairs of smaller spines. Male metatibiae arcuate, Anterior
face of metatibiae concolorous with posterior face; external apical tubercle present.

Length (mm) – Range: 23.5 - 29.0, X = 26.0; males: 27.2 and 29.0; female range: 23.5 -

26.7, X = 25.0.

Width at pronotum (mm) – Range: 7.6 - 8.9, X = 8.3; males: 8.5 and 8.9; female range:

7.6 - 8.7, X = 8.1.

Width at connexiva (mm) – Range: 8.2 - 8.9, X = 8.6; males: 8.6 and 8.9; female range:

8.2 - 8.8, X = 8.5.
Diagnosis – This species is readily distinguished by possessing and a pronotum margined
anteriorly with a red semilunar band that extends to the lateral angles of the pronotum and
is over 23 mm in length. The brownish-black to black color and bicolored connexiva may
cause confusion of P. rubronotatus with P. rubropictus, but the two are readily separated
by the presence of the red semilunar band (Fig. 94, 96) and a trigonally-shaped area
surrounding the abdominal segment spiracle IV in males, both of which P. rubropictus
lacks. P. rubronotatus may also be confused with P. vates, because both are approximate
in length and width and are brownish-black to black in color, but P. vates lacks red
ventroabdominal markings and the red pronotal semilunar band. Moreover, antennal
segment IV and tarsi are never accented with color in P. rubronotatus.

57
Distribution – (Fig. 22) Belize (UMC); Guatemala; Mexico: Oaxaca, Queretaro
(NMNH) San Luis Potosi, Tamaulipas, Veracruz.
Piezogaster rubropictus (Montandon)

(Fig. 98-101)
Archimerus rubropictus Montandon 1897; p. 248; Ecuador.

Piezogaster rubropictus: O'Shea 1980; p. 214.
Body color brownish-black to black with reddish-orange to orange accents, covered
with minute hairs. Postocular area concolorous with head, sometimes tuberculate.
Antennal segment I longest, segment II, IV subequal, segment III shortest; all segments
concolorous. Trace of laterally projecting jugal shelf present beneath base of antennal
segment I in males; not present in females. Beak segments I, II, IV subequal in length,
segment III shortest. Head region just anterior to pronotal collar extending anteriorly to
posterior ocellar margin reddish-orange to orange in color; anterior pronotal collar
punctate. Anterior pronotal margins slightly tuberculate, callar region impunctate,
sparsely haired with minute tubercles. Remainder of pronotum strongly tuberculate, with
four reddish-orange to orange longitudinal stripes; lateral-most stripes running length of
pronotal margin; interior stripes not reaching posterior margin of pronotum, running
parallel to each other. Pronotal expansions lacking. Posterior pronotal angles lacking.
Scutellum reddish-orange to orange with dark brown to black margin and a medial dark-
brown to black stripe dissipating before reaching the posterior tip. Corium mottled with
reddish-orange to orange and brownish-black to black. Membrane brownish-black,
venation with little or no anastomosis. Pronotal, abdominal widths subequal in males;

58
abdomen slightly wider than pronotum in females. Connexival segments with minute
hairs, tubercles lacking; segments II-VI bicolored; anterior orange to reddish-orange,
posterior brownish-black to black; segment VII entirely orange to reddish-orange.
Posterolateral spur present on segments IV-VI, sometimes III. Abdomen lacking ventral
markings. Genital capsule rim notched; protuberances lateral to each side of notch
margined with minute hairs. In posterior view, genital capsule medially rugose, dorsally
depressed. Propleural acetabular suture not keeled. Red to reddish-orange mark present
on each thoracic pleura. Posterior metapleural margin straight. Area surrounding
and mesofemora with ventrodistal pair of minute ultimate spines; penultimate spines much
larger. Interior face of metatrochanter with flattened spine in males and interior ventral
face of metafemora with three spines extending from distal end halfway down the length;
two pairs of ventrodistal spines in series in females. Anterior face of metatibiae
concolorous with posterior face; male metatibiae arcuate, external apical tubercle lacking.


range: 20.3 - 22.2, X = 21.0.


female range: 6.7 - 8.1, X = 7.3.


female range: 7.4 – 8.8, X = 7.9.
Diagnosis – This species is similar to P. camposi because of the reddish-orange and
brownish-black to black coloration. However, P. rubropictus lacks the prominent lateral

59
expansions of P. camposi. P. rubropictus also lacks the brownish-black to black macula
on each corium. Additionally, the head of P. camposi is always reddish-orange to red,
while the head of P. rubropictus is brownish-black to black, and P. camposi has a pale
orange colored metathoracic scent gland area, as opposed to the darker scent gland of P.
rubropictus.
Distribution – (Fig. 23) Bolivia (NMNH); Ecuador; Peru.
Piezogaster scutellaris Stål

(Fig. 102-105)
Piezogaster scutellaris Stål 1862; p. 292; Mexico.

Archimerus scutellaris: Stål 1867; p. 538.
Archimerus maculifer Walker 1871. Synonymized by Distant (1881).
Piezogaster scutellaris: O'Shea 1980; p. 214.
Body color medium brown to pale brown; extremities usually tinged with red or
orange. Covered in short, often decumbent white hairs. Head with short white hairs,
tuberculate. Postocular area darker than remainder of head with lighter tubercles.
Antennal segment I longest, segments II, IV subequal, III shortest; color of ultimate
antennal segment variable. Laterally projecting jugal shelf present beneath base of
antennal segment I. Beak segments I, II, IV subequal, III shortest. Pronotal collar
punctate, often tuberculate. Anterior margin of pronotum with small tubercles extending
to lateral angles. Callar region with white decumbent hairs, impunctate; remainder of
surface punctate, often tuberculate. Humeral angles markedly widened (Fig. 140).
Posterior pronotal angles present. Scutellum usually yellowish in color; punctate, darkly
margined. Corium concolorous with body, membrane often slightly darker, venation
60
slightly anastomosing. Pronotum equal in width to abdomen or slightly wider. Anterior
third to half of connexival segments yellowish in color, tubercles lacking, impunctate, hairs
sometimes apparently velvety. Posterolateral connexival spurs minute to lacking. Venter
of abdominal segments III-VII each with a pair of minute yellowish spots. Genital capsule
rim entire, curving slightly; margined with minute hairs. Genital capsule seen from
posterior view rugose; minute punctations barely visible. Propleural acetabular suture
lacking keel. Posterior metapleural margin sinuous. Area surrounding abdominal segment
IV spiracle not raised as trigonal area. Metacoxal knob lacking. Pro- and mesofemora
with pale yellow tubercles; ventrodistally armed with pair of small darker spines followed
by pair of much larger darker spines, followed by 3-4 pairs of pale tubercles; distal-most
pair of tubercles sometimes spinose. Metafemora strongly crassate in males, armed with
large widened spines or spinose tubercles, pale yellow in color, similar in arrangement to
those of pro- and mesofemora; ventrodistal most spines fused as one spine with double-
point. Anterior face of metatibiae concolorous with posterior face; metatibiae slightly
arcuate; external apical tubercle present in males.


range: 15.5 - 17.6, X = 16.9.


female range: 5.3 - 6.2, X = 5.9.


female range: 4.9 - 6.5, X = 5.5.
Diagnosis – This species appears to be a smaller version of P. chiriquinus because they

61
both share an elongate-looking body form and bicolored connexiva. However, P.
scutellaris lacks the lateral expansions of P. chiriquinus (Fig.134), and instead has
markedly widened humeral angles (Fig. 140). P. scutellaris also has more incrassate
metafemora than P. chiriquinus, and the metafemoral tubercles of P. scutellaris are pale
yellow; the metafemoral tubercles of P. chiriquinus are never pale yellow.
Distribution – (Fig. 24) Belize (CAS); Costa Rica (UCB); Guatemala (UMC);
Honduras (NMNH); Mexico: Chiapas, Colima, Guerrero, Hidalgo, Jalisco, Michoacan,
Morelos, Nuevo Leon, Oaxaca, Puebla, Queretaro, San Luis Potosi, Tamaulipas
(TAMU), Veracruz, Yucatan; Panama (TAMU); Nicaragua.
Notes – I examined one homoeotype from UMC designated and labeled as such by
Thomas Yonke. Schaefer & O'Shea (1979) noted Oryza sativa as a host plant.
Piezogaster spurcus (Stål)

(Fig. 106-109)
Capaneus spurcus Stål 1862; p. 291; Mexico.

Piezogaster spurcus: O'Shea 1980; p. 214.
Body color light brown to cinnamon-brown, covered entirely in short, white hairs and
tubercles, except the anterior surface of pronotum, scutellum, and hemelytra, giving the
entire body a tomentose appearance. Postocular area darkest part of head, strongly
tuberculate. Antennal segment II longest, segments I, IV slightly shorter than segment I,
subequal; segment III shortest; all segments concolorous. Laterally projecting jugal shelf
present beneath base of antennal segment I. Beak segments I, II, IV subequal in length,
segment III shortest. Anterior pronotal collar turberculate, punctate. Anterior margins of
62
pronotum tuberculate; callar region with short, erect hairs, tuberculate, punctations
lacking; remainder of pronotum heavily punctate; lateral expansions lacking. Posterior
pronotal angles lacking. Scutellum heavily punctate, sometimes darkly margined on
posterior sides. Corium concolorous with body, membrane apparently darker than
remainder of body with venation anastomosing. Abdomen slightly wider than pronotum.
Connexiva unicolorous, tuberculate. Posterolateral connexival spur present on abdominal
segments III-VII, sometimes II; spur noticeably larger on segments V-VII. Abdomen
lacking ventral markings. Genital capsule rim entire, straight; margined with hairs.
Genital capsule in posterior view medially rugose; covered in punctations and minute
hairs. Propleural acetabular suture lacking keel. Posterior metapleural margin sinuous.
Area surrounding abdominal segment IV spiracle not raised as trigonal area. Prominent
knob present, arising laterally from metacoxa, knob lacking hook. Ventrodistal of
profemora with 3-4 pairs of spines becoming smaller and paler as the series extends
proximally; distal-most spine significantly larger. Ventrodistal of mesofemora with 4-5
pairs of spines becoming smaller and paler as the series extends distally; distal-most spine
significantly larger. Dorsal side of metafemora covered in large tubercles, sometimes in
rows running lengthwise; venter of metafemora with five pairs of darker spines; in males,
most proximal interior spine significantly larger. Venter of metatibiae with three
significantly larger, darker spines; spine arrangement more prominent in males. Interior
face of entire hind leg always darker than exterior face. Male metatbiae arcuate; external
apical tubercle lacking.

63


range: 17.0 - 19.6, X = 18.0.


female range: 6.2 - 7.5, X = 6.7.


female range: 8.3 - 9.6, X = 8.8.
Diagnosis – This species is one of the few that has tubercles covering the connexiva. It is
also recognized by the uniquely larger spur arising from connexival segments V, VI and
VII, and a covering of short white hairs. P. spurcus sometimes may be confused with P.
yonkei. However, P. yonkei has more of a rose or sanguine color, does not appear nearly
as tomentose, has defined posterior pronotal angles, and has two distinct knobs on its
genital capsule. Additionally, P. spurcus always has three or more spines present on the
venter of its metafemora.
Distribution – (Fig. 25) Costa Rica (UCB); Guatemala; Mexico: Campeche, Colima,
Durango, Estado De Mexico, Guerrero, Jalisco, Morelos, Nayarit, Queretaro (UCB),
San Luis Potosi, Sonora, Yucatan Zacatecas (TAMU); United States: Arizona (UMC).
Notes – I examined two homoeotypes from UMC designated and labeled as such by
Thomas Yonke.
Piezogaster tetricus (Stål)

(Fig. 110-113)
Capaneus tetricus Stål, 1862; p. 291; Mexico.

Archimerus muticus Walker 1871. Synonymized by Distant (1881).
Piezogaster tetricus: O'Shea 1980; p. 214.
64
Body color pale to dark brown, covered with minute hairs. Head covered with
minute tubercles. Postocular area often darker than remainder of head, always
tuberculate. Antennal segment I longest, segments II, IV subequal in length, segment III
shortest; segment IV sometimes apparently paler than other segments. Laterally
projecting jugal shelf present beneath base of antennal segment I. Beak segments I, II, IV
subequal, segment III shortest. Anterior pronotal collar laterally punctate. Anterior
margins of pronotum with minute tubercles, callar region impunctate, sometimes sparsely
tuberculate; remainder of pronotum punctate; lacking lateral expansions. Posterior
pronotal angles lacking. Scutellum sometimes paler than remainder of body, rugosely
punctate, sometimes darkly margined. Corium usually concolorous with body.
Membrane of hemelytra apparently darker than than remainder of body; venation with
little or no anastomosis. Pronotum wider than abdomen, sometimes subequal in females.
Connexival segments with minute hairs, yellowish at anterolateral corner, apparently
bicolored; tubercles lacking. Minute posterolateral connexival spur on abdominal
segments III-VI, sometimes II. Pair of minute dorsoabdominal spots often present on
each visible abdominal segment. Genital capsule rim entire, sometimes slightly dorsally
depressed, margined with minute hairs. In posterior view, genital capsule medially rugose,
punctate. Propleural acetabular suture not keeled. Posterior metapleural margin straight.
In males, area surrounding abdominal segment IV spiracle raised, trigonal. Metacoxal
knob lacking. Pro- and mesofemora with ventrodistal pair of minute ultimate spines;
penultimate spines much larger, preceded proximally by 2-3 pairs of sometimes spinose
tubercles. In males, metafemora slightly arcuate, with same ventral pattern as pro- and
mesofemora, but most distal spines often fused into a single double-pointed spine; females
65
with up to five pairs of spines or spinose tubercles proximally following double-pointed
spine. Anterior face of metatibiae concolorous with posterior face; metatibiae vaguely
arcuate, males with external apical tubercle.


range: 19.0 - 22.6, X = 20.7.


female range: 6.4 - 7.6, X = 6.9.


female range: 6.0 - 7.1, X = 6.6.
Diagnosis – P. tetricus is similar morphologically to P. odiosus and P. basilicus, all three
having a pronotum wider or subequal to the abdominal width (Fig. 142). However, P.
tetricus is usually paler than P. odiosus and has contrasting bicolored connexival segments
that are easily seen with naked eye. P. basilicus is longer, 24 - 25 mm, and is
proportionately narrower than P. tetricus in the anterior abdominal region as well.
Distribution – (Fig. 26) Costa Rica (TAMU); El Salvador (UMC); Guatemala;
Honduras (NMNH); Mexico: Chiapas, Colima (TAMU), Durango, Estado De Mexico,
Guerrero, Hidalgo, Jalisco, Michoacan, Morelos, Nayarit (UMC), Nuevo Leon, Puebla
(NMNH), Oaxaca, Quintana Roo, San Luis Potosi, Tabasco, Tamaulipas (TAMU)
Veracruz, Yucatan.
Notes – Brailovsky & Barrera (1984) stated that the main difference between P. basilicus
and P. tetricus is that P. tetricus is not longer than 21 mm, whereas P. basilicus is
between 24-25 mm. However, I found specimens of what was apparently P. tetricus that
66
ranged from 19 - 23.8 mm in length, but did not exceed 24 mm. Moreover, the ranges of
these two species overlap in several places. Lack of an adequate number of identified P.
basilicus specimens prevents me from drawing absolute conclusions, because the main
character separating the two species is a difference in size.
I examined four homoeotypes from UMC designated and labeled as such by
Thomas Yonke.
Piezogaster thoracicus (Distant)

(Fig. 124-125)
Archimerus thoracicus Distant 1881; p. 114; Guatemala.

Piezogaster thoracicus: O'Shea 1980; p. 214.
This species was described by Distant (1881) as having a fuscous body with a paler,
concolorous underside and pronotal angles that are dilated with a subacutely pointed
apex, and finely denticulate on lateral margins. He also described this species as having a
transversely and irregularly striate scutellum and incrassate metafemora, especially
pronounced in the males. Judging from Distant's descriptions and illustrations, this
species appears similar to P. chiriquinus but has wider, more laterally produced pronotal
expansions which terminate in subacute points (Fig. 130).
Distant's original description cites seven males and eight females, but I have not
personally seen any specimens for this species. The only other reference to P. thoracicus I
found in my literature review was its combination with Piezogaster by O'Shea (1980).
However, Distant noted the distinctly dilated pronotal angles were enough to discern this
species as unique.
67
Measurements – From Distant (1881). Males: n = 7; females: n = 8.
Length (mm) – Range: 20 – 22.
Width at pronotum (mm) – Range: 9 – 10.
Distribution – (Fig. 27) Guatemala.
Piezogaster vates (Stål)
(Fig. 114-117)
Capaneus vates Stål, 1862; p. 290. Mexico.

Piezogaster vates: O'Shea 1980; p. 214.
Body more than 23 mm in length, color brownish-black to black, covered with
minute hairs. Head sparsely tuberculate to lacking. Postocular area as dark as remainder
of head, with paler tubercles. Antennal segment I longest, segments II, IV subequal in
length, segment III shortest. Ultimate antennal segment orangish-red to orange in color.
segments I, II, IV subequal, segment III shortest. Lateral sides of anterior pronotal collar
punctate. Anterior margins of pronotum with minute tubercles, callar region sometimes
tuberculate, impunctate. Remainder of pronotum punctate. Lateral pronotal expansions
lacking. Posterior pronotal angles lacking. Scutellum rugosely punctate. Hemelytra
concolorous with body, membrane venation with little or no anastomosis. In males,
pronotum wider than abdomen, width subequal in females. Connexiva with minute hairs,
unicolorous, tubercles lacking. Minute, posterolateral spur present on segments IV-VI,
sometimes III. Abdomen lacking ventral markings. Genital capsule rim entire, sometimes
68
depressed dorsally. Viewed posteriorly, genital capsule medially rugose, punctate.
Propleural acetabular suture not keeled. Posterior metapleural margin straight. In males,
area surrounding abdominal segment IV spiracle forming a raised trigonal area.
Metacoxal knob lacking. Pro- and mesofemora with ventrodistal pair of minute ultimate
spines; penultimate spines much larger, preceded proximally by 2-3 pairs of sometimes
spinose tubercles. Dorsal and lateral surfaces of metafemora marked sparsely with
spinose tubercles. In males, metafemora slightly arcuate, with same ventral pattern as
pro- and mesofemora, but distal-most spines often fused into a single double-pointed
spine; females with up to five pairs of spines or spinose tubercles proximally following
double-pointed spine. Inner, outer face of metatibiae concolorous. Male metatibiae
arcuate with external apical tubercle. All tarsi same orangish-red to orange color as
antennal segment IV.
Measurements – Males: n = 2; female: n = 1.
Length (mm) – Males: 26.4 and 27.9; female: 25.9.
Width at pronotum (mm) – Males: 8.4 and 9.1; female: 8.8.
Width at connexiva (mm) – Males: 7.7 and 8.2; female: 8.6.
Diagnosis – P. vates is morphologically similar to P. tetricus, P. rubronotatus, and P.
odiosus, all having a pronotum that is narrower or subequal to the width of the abdomen.
However, by my measurements, and those of Brailovsky & Barrera (1984), P. vates is
longer than P. odiosus (more than 23 mm whereas P. odiosus is shorter than 23 mm).
The ultimate antennal segments and tarsi of P. vates are usually orangish-red to orange in
color, similar to P. alienatus. Aside from usually being much longer and wider than P.
tetricus, P. vates also lacks the pale anterolateral spot on each connexival segment found
69
in P. tetricus. Although almost equal in length and both pronotal and abdominal widths,
P. vates lacks the red to reddish-orange semilunar pronotal band of P. rubronotatus, and
has orangish-red to orange ultimate antennal and tarsal segments, which P. rubronotatus
lacks.
Distribution – (Fig. 28) Mexico: Jalisco, Morelos, Nayarit, Oaxaca, Quintana Roo,
Veracruz.
Notes – P. vates appears to be misrepresented in many collections by a misidentification of
the darker variant of P. odiosus. Stål's original description of P. vates makes note of its
significantly larger size, and I concur that it is the more important character when
separating P. vates and P. odiosus.
Piezogaster yonkei sp. nov.

(Fig. 118-121)
(Description of holotype)
Body color rose or sanguine, covered with short white hairs. Head tuberculate.
Postocular area darker than remainder of head, tuberculate. Antennal segment II longest,
segments I, IV slightly shorter than segment I, subequal; segment III shortest; all
segments concolorous. Laterally projecting jugal shelf present beneath base of antennal
segment I. Beak segments I, II, IV subequal in length, III shortest. Lateral areas of
pronotal collar strongly punctate. Anterior margin of pronotum tuberculate, remaining
margin not tuberculate. Callar region somewhat tuberculate, with hairs, impunctate;
remainder of pronotum strongly punctate; lateral expansions lacking. Posterior pronotal
angles present. Scutellum punctate, slightly rugose. Hemelytra roughly concolorous with
70
body, venation of membrane anastomosing. Abdomen wider than pronotum. Connexiva
with minute hairs, tuberculate. Lateral margins of connexival segments with alternating
dark brown to black and pale yellow spots. Posterolateral spur present on connexival
segments II-VII, Abdomen lacking ventral markings. Genital capsule rim entire, straight,
margined with minute hairs. Genital capsule in posterior view entirely punctate with
minute hairs, medially rugose; armed with two protuberances just underneath and lateral
to posterodorsal margin. Propleural acetabular suture smooth. Posterior metathoracic
margin sinuous. Pro- and mesofemora with ventrodistal pair of minute ultimate spines;
penultimate spines much larger. Area surrounding abdominal segment IV spiracle not
raised as trigonal area. Metacoxa with prominent lateral knob, knob lacking distal hook.
Metafemora with distinct raised medial region adorned with a single large spine. Anterior
face of metatibiae darker than posterior face. Metatibiae arcuate; external apical tubercle
lacking.
Holotype: male (Fig. 118-119). Length: 19.9 mm; width at pronotum: 8.0 mm; width at
connexiva: 9.4 mm. Mexico: Durango. July 6, 1952. Collected by J.D. Lattin. Holotype
designated from and deposited in the entomology collection of University of California,
Berkeley.
Measurements of all specimens – Males: n = 13; females: n = 10.


range: 17.2 - 20.9, X = 19.2.

Width at pronotum (mm) - Range: 6.2 - 8.2, X = 7.4; male range: 6.2 - 8.2, X = 7.3;

female range: 6.4 - 8.0, X = 7.5.

71

female range: 7.7 - 9.8, X = 8.9.
Diagnosis – This newly described species probably is confused with P. spurcus in many
collections. However upon examination, it is quite evident that the rose or sanguine color
of the body of P. yonkei is all that is needed to separate the two. Although diagnosis for
the female is mostly a difference of color, P. yonkei has posterior pronotal angles which
P. spurcus lacks. Additionally, P. yonkei males have a single ventral metafemoral spine
and two protuberances on its genital capsule, whereas P. spurcus males have at least three
metafemoral tubercles and lack the genital capsule protuberances.
Distribution – (Fig. 29) United States: Arizona (TAMU); Mexico: Chihuahua
(AMNH), Distrito Federal (AMNH), Durango (UCB), Michoacan (UCB), Sinaloa
(AMNH).
Notes – This species is named in honor Thomas Yonke, whose research and labeling of
homoeotype specimens was invaluable to my research. Moreover, among the specimens
of P. yonkei I received, one was identified a "sp. nov." by Yonke, but had never been
followed up by him. For these reasons, I find it only fitting that this specimen bear his
name.
Female characters from paratypes. Female metafemora lacking distinct raised
medial region adorned with a single large spine. Female metatibia not arcuate.
Posterolateral spur on connexival segment II sometimes lacking. Paratypes deposited in
AMNH, SMHP, TAMU, and UCB.

72
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north of Mexico. Part II. Entomologica Americana. 21(2): 41-122.
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U.S. Geologic and Geographic Survey of the Terrain (2nd Ser.). 5: 296-297.
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excepting the Aphididae, Coccidae and Aleurodidae. Berkeley: University of
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of the British Museum. Part IV. London: British Museum. 1-211.
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Yonke, T. R. and J. T. Medler. 1969a. Biology of the Coreidae in Wisconsin.

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477-480.
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
Figure 32-33: P. achilles; male – dorsal and lateral views. From NMNH collection.
Figure 34-35: P. achilles; female – dorsal and lateral views. From NMNH collection.
93
Figure 36-37: P. acuminatus holotype; male – dorsal and lateral views. From CMNH collection.
94
Figure 38-39: P. alienatus paratype; male – dorsal and lateral views. From TAMU collection.
Figure 40-41: P. alienatus holotype; female – dorsal and lateral views. From TAMU collection.
95
Figure 42-43: P. alternatus; male – dorsal and lateral views. From SMHP collection.
Figure 44-45: P. alternatus; female – dorsal and lateral views. From TAMU collection.
96
Figure 46-47: P. auriculatus; male – dorsal and lateral views. From CAS collection.
Figure 48-49: P. auriculatus; female – dorsal and lateral views. From CMNH collection.
97
Figure 50-51: P. basilicus; male – dorsal and lateral views. From TAMU collection.
Figure 52-53: P. basilicus; female – dorsal and lateral views. From UCB collection.
98
Figure 54-55: P. calcarator; male – dorsal and lateral views. From UMC collection.
Figure 56-57: P. calcarator; female – dorsal and lateral views. From CMNH collection.
99
Figure 58-59: P. camposi; male – dorsal and lateral views. From Distant collection at BMNH.
Figure 60-61: P. camposi; female – dorsal and lateral views. From SMEK collection.
100
Figure 62-63: P. chiriquinus; male – dorsal and lateral views. From NMNH collection.
Figure 64-65: P. chiriquinus; female – dorsal and lateral views. From NMNH collection.
101
Figure 66-67: P. chontalensis; male – dorsal and lateral views. From Distant collection at BMNH.
Figure 68-69: P. chontalensis; female – dorsal and lateral views. From Distant collection at BMNH.
102
Figure 70-71: P. dilatatus; male – dorsal and lateral views. From CMNH collection.
Figure 72-73: P. dilatatus; female – dorsal and lateral views. From NMNH collection.
103
Figure 74-75: P. humerosus; female – dorsal and lateral views. From NMNH collection.
104
Figure 76-77: P. indecorus; male – dorsal and lateral views. From NMNH collection.
Figure 78-79: P. indecorus; female – dorsal and lateral views. From NMNH collection.
105
Figure 80-81: P. obscuratus; male – dorsal and lateral views. From NMNH collection.
Figure 82-83: P. obscuratus; female – dorsal and lateral views. From NMNH collection.
106
Figure 84-85: P. odiosus; male – dorsal and lateral views. From UCB collection.
Figure 86-87: P. odiosus; female – dorsal and lateral views. From UMC collection.
107
Figure 88-89: P. odiosus, lighter colored variant; male – dorsal and lateral views. From UMC collection.
108
Figure 90-91: P. reclusus; male – dorsal and lateral views. From NMNH collection.
Figure 92-93: P. reclusus; female – dorsal and lateral views. From UMC collection.
109
Figure 94-95: P. rubronotatus; male – dorsal and lateral views. From NMNH collection.
Figure 96-97: P. rubronotatus; female – dorsal and lateral views. From TAMU collection.
110
Figure 98-99: P. rubropictus; male – dorsal and lateral views. From NMNH collection.
Figure 100-101: P. rubropictus; female – dorsal and lateral views. From NMNH collection.
111
Figure 102-103: P. scutellaris; male – dorsal and lateral views. From FMNH collection.
Figure 104-105: P. scutellaris; female – dorsal and lateral views. From TAMU collection.
112
Figure 106-107: P. spurcus; male – dorsal and lateral views. From UCB collection.
Figure 108-109: P. spurcus; female – dorsal and lateral views. From SMHP collection.
113
Figure 110-111: P. tetricus; male – dorsal and lateral views. From UMC collection.
Figure 112-113: P. tetricus; female – dorsal and lateral views. From UMC collection.
114
Figure 114-115: P. vates; male – dorsal and lateral views. From NMNH collection.
Figure 116-117: P. vates; female – dorsal and lateral views. From NMNH collection.
115
Figure 118-119: P. yonkei holotype; male – dorsal and lateral views. From UCB collection.
Figure 120-212: P. yonkei paratype; female – dorsal and lateral views. From AMNH collection.
116
Figure 122-125 from Distant (1880-1893).
Fig. 122: P. loricata; Fig. 123: P. multispinus; Fig. 124-125: P. thoracicus, male and female,
respectively.
117
Figure 126-133: Piezogaster pronota showing lateral expansions. Fig. 126: P. achillelus, Fig. 127: P.
reclusus, Fig. 128: P. achilles, Fig. 129: P. chontalensis, Fig. 130: P. thoracicus, Fig. 131: P. humerosus,
Fig. 132: P. acuminatus, Fig. 133: P. obscuratus. Fig. 126-129 reproduced from Brailovsky & Barrera
(2000). Fig. 130 reproduced from Distant (1880-1893).
118
Figure 134-140: Piezogaster pronota. Fig. 134: P. chiriquinus, Fig. 135: P. auriculatus. Figures 136-
139: Piezogaster pronota lacking lateral expansions. Fig. 136: P. alienatus, Fig. 137: P. odiosus, Fig.
138: P. yonkei, Fig. 139: P. calcarator. Figure 140: P. scutellaris pronotum with widened lateral angles.
Figure 141: genital capsule of P. congruus, reproduced from Brailovsky & Barrera (1983).
119
Figure 142: Body of P. odiosus, abdomen narrower or nearly as wide as pronotum.
Figure 143: Body of P. alienatus, abdomen wider than pronotum.

120
Fig. 144
Figure 144: Areas measured on specimens – a. Total length from tip of tylus to the end of the abdomen.
b. Width of the pronotum at the widest point. c. Width of the abdomen at the widest point.
121
Figure 145: Relative lengths for all species (in millimeters).

† Denotes measurements from original description.

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A REVISION OF THE GENUS PIEZOGASTER AMYOT & SERVILLE

(HETEROPTERA: COREIDAE: NEMATOPODINI)

AND THE DESCRIPTION OF

TWO NEW SPECIES

A Thesis Presented to the Graduate Faculty

of the Fort Hays State University in

Partial Fulfillment of the Requirements for

the Degree of Master of Science

B.S., Fort Hays State University

Description of the genus Piezogaster Amyot & Serville..................................................8

Diagnostic key to known species....................................................................................10

Species descriptions (alphabetical listing) –

Piezogaster achillelus Brailovsky & Barrera......................................................13

Distributional data (Fig. 1-31)...........................................................................76

Images of specimens (Fig. 32-121)....................................................................92

Illustrations (Fig. 122-144)..............................................................................116

Relative lengths for all species (Fig. 145).........................................................121

Minnesota, St. Paul.

(Heteroptera: Coreidae: Nematopodini)

and the Description of

Two New Species

The New World Heteropteran genus Piezogaster is revised with in-depth

alternatus (Say) is resurrected from synonymy under Piezogaster calcarator (Fabricius),

after examination of homoeotype specimens, ranges and examination of the descriptions of

both species in the literature. Piezogaster ashmeadi (Montandon) is synonymized with P.

complete lack of identified specimens of P. ashmeadi. Piezogaster herrichi (Blöte) is

synonymized with Piezogaster indecorus (Walker), because of the vagueness of the

original description of P. herrichi, the reliance on an inconsistent diagnostic character for

separation, and sympatric ranges of both species. Piezogaster humeralis (Distant) is

synonymized with Piezogaster camposi (Montandon), based on prior research by O'Shea

Distant's collection at the British Museum identified as P. humeralis that is identical in

is synonymized with Piezogaster auriculatus (Stål) after examination of original

both species, using specimens identified by Brailovsky.

species. Distributions examined using Geographic Information System software clarified

questions concerning range overlap of P. alternatus and P. calcarator as well as P.

heteropterans, and, historically, has exhibited much systematic confusion. My research

groundwork for further research on its members.

that the genus Archimerus originally was established by Burmeister to be a replacement

for the preoccupied Pachymeria Laporte (1833). Archimerus Burmeister and

Archimerus as a valid nematopodine genus. The next available genus is Piezogaster, so

extensive heteropteran catalog (1870); Distant's catalog of Central American Heteroptera

My objective is to provide a species-level revision of the genus Piezogaster Amyot &

I conducted an extensive literature search on Piezogaster, and compiled a list of

acuminatus Brailovsky, and homoeotypes, specimens compared directly with the

holotype, for Piezogaster auriculatus (Stål), Piezogaster calcarator (Fabricius),

foundation to add inaccessible species. This was accomplished by incorporating

discovered distributions through my research. Distributional data are given in each

species description, as well as graphically represented (Figures 1-31). New distributions

(American Museum of Natural History, NewYork), BMNH (British Museum of Natural

History, London), CAS (California Academy of Sciences, San Francisco), CMNH

(Carnegie Museum of Natural History, Pittsburgh), CU (Cornell University, Ithaca),

FMNH (Field Museum of Natural History, Chicago), NMNH (National Museum of

Natural History, Washington, DC), SMEK (Snow Museum of Entomology, University of

(University of California, Berkeley), UMAA (University of Michigan, Ann Arbor), UMC

(University of Missoiri, Colombia), UMSP (University of Minnesota, St. Paul).

Coolpix 900 series digital camera affixed to a stationary platform.

From my research, I described two new species of Piezogaster, one exclusive to

species, updated and expanded distributional data, and produced a diagnostic

dichotomous key to species.

Piezogaster alternatus (Say) is resurrected from synonymy under P. calcarator after

examination of homoeotype specimens, ranges and examination of the descriptions of both

species in the literature. I synonymize Piezogaster ashmeadi (Montandon) with P.

complete lack of identified specimens of P. ashmeadi. Piezogaster herrichi (Blöte) is