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

ii

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.

iii

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.19801999), 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 reddishorange. 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 1' 2(1) 2'

Pronotal expansions present (Fig.126-135) or humeral angles markedly widened (Fig. 140)...........................................................................................2 Pronotal expansions lacking, pronotal angles not widened (Fig.136-139).........17 Pronotal margin strongly dentate-serrate, femora extremely multispinose (Fig. 123).....................................................................................P. multispinus (Stål)* Pronotal margin not strongly dentate-serrate; may be tuberculate or weakly serrate (Fig. 126-140). If femoral spines present, not abundant.......................3 Pronotum covered in strong, rounded, reddish-brown to black tubercles, more numerous on rising lateral expansions (Fig. 131) ..............P. humerosus (Distant) Pronotum lacking strong, rounded, reddish-brown to black tubercles, or if present, lacking on expansions......................................................................................4 Pronotum expanded laterally, terminating in subacute points (Fig. 130) ........... ........................................................................................P. thoracicus (Distant)* Pronotum lacking lateral expansions, or if laterally expanded, lacking subacute points..............................................................................................................5 Laterally projecting jugal shelf present beneath base of antennal segment I.......6 Lacking laterally projecting jugal shelf beneath base of antennal segment I ......11 Antennal segment IV concolorous with remainder of segments........................7 Antennal segment IV not concolorous with remainder of segments..................10 With a pale yellow sternal fascia on each thoracic pleura (Fig. 81, 83).............. ........................................................................................................................8 Sternal fascia lacking.......................................................................................9 Body black; with 4 pairs of pale yellow abdominal discoid areas running ......... in series................................................................................P. loricata (Distant)* Body medium to dark brown, never black; abdominal discoid areas lacking or reduced to minute or obscure spots.............................P. obscuratus (Montandon) Pronotal expansions rounded, earlike (Fig. 135); abdomen wider than pronotum ..............................................................................................P. auriculatus (Stål) Pronotal expansions not rounded, terminating in a slightly anteriorly directed acute angle (Fig. 129); pronotum wider than abdomen ............P. chontalensis (Distant)

3(2) 3'

4(3) 4'

5(4) 5' 6(5) 6' 7(6) 7' 8(7) 8'

9(8) 9'

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 proand 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. Width at pronotum (mm) – Range: 6.1 - 7.5, X = 6.8; male range: 6.3 - 7.5, X = 6.9; female range: 6.1 - 7.4, X = 6.6. Width at connexiva (mm) – Range: 7.2 - 9.3, X = 8.0; male range: 7.3 - 9.3, X = 8.2; 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.
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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. Measurements – Males: n = 11; females: n = 10. Length (mm) – Range: 20.0 - 25.5, X = 22.2; male range: 21.6 - 25.5, X = 23.1; female 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 reddishorange 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
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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. Laterally projecting jugal shelf present beneath base of antennal segment I. Beak 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. Measurements – Males: n = 1; females: n = 1. 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. Length (mm) – Male: 18.3; female: 18.6. Width at pronotum (mm) – Male: 7.9; female: 7.6. 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. Proand 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 Measurements – Males: n = 10; females: n = 10. 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. Width at pronotum (mm) – Range: 5.9 - 7.4, X = 6.8; male range: 5.9 - 7.4, X = 6.9; female range: 6.3 - 7.0, X = 6.7. Width at connexiva (mm) – Range: 7.1 - 9.6, X = 8.8; male range: 7.1 - 9.4, X = 8.7; 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.
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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 reddishorange 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 abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob lacking. Proand 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. Measurements – Males: n = 1; females: n = 4. 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
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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. Measurements – Males: n = 10; females: n = 10. 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. Width at pronotum (mm) – Range: 6.2 - 8.3, X = 7.4; male range: 6.2 - 8.3, X = 7.5; female range: 7.0 - 7.9, X = 7.4. Width at connexiva (mm) – Range: 5.3 - 8.1, X = 6.7; male range: 5.3 - 7.1, X = 6.2; 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.
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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 surrounding abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob

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. Measurements – Males: n = 2; females: n = 2. 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.
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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. Length (mm) – Male: 20.8; female: 18.7. Width at pronotum (mm) – Male: 6.3; female: 5.6. 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 abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob lacking. Pro-

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. Measurements – Males: n = 5; females: n = 4. Length (mm) – Range: 17.1 - 20.5, X = 18.6; male range: 17.1 - 19.3, X = 18.0; female range: 18.6 - 20.5, X = 21.4. Width at pronotum (mm) – Range: 6.6 - 8.0, X = 7.3; male range: 6.6 - 8.0, X = 7.2; female range: 7.0 - 8.0, X = 7.5. Width at connexiva (mm) – Range: 7.3 - 11.2, X = 9.4; male range: 8.5 - 9.5, X = 9.1; 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.
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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 surrounding abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob 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. Measurements – Males: n = 10; females: n = 10. Length (mm) – Range: 17.6 - 22.5, X = 19.7; male range: 19.1 - 22.5, X=20.2; female range: 17.6 - 21.6, X = 19.3. Width at pronotum (mm) – Range: 6.3 - 8.4, X = 7.3; male range: 6.8 - 8.4, X = 7.3; female range: 6.3 - 8.0, X = 7.2. Width at connexiva (mm) – Range: 6.7 - 10.0, X = 8.1; male range: 7.4 - 8.7, X = 8.3; 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
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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. Measurements – Males: n = 9; females: n = 4. Length (mm) – Range: 16.8 - 25.5, X=20.0; male range: 16.8 - 25.5, X=19.8; female range: 18.6 - 22.5, X = 20.6. Width at pronotum (mm) – Range: 6.7 - 11.9, X = 8.7; male range: 6.7 - 11.9, X = 8.5; female range: 7.7 - 10.3, X = 9.1. Width at connexiva (mm) – Range: 6.8 - 11.5, X = 9.1; male range: 6.8 - 11.5, X = 8.7; 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
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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. Measurements – Males: n = 10; females: n = 10. Length (mm) – Range: 16.7 - 22.8, X = 20.4; male range: 19.0 - 22.7, X=20.8; female range: 16.7 - 22.8, X = 19.9. Width at pronotum (mm) – Range: 5.3 - 7.3, X = 6.7; male range: 6.2 - 7.3, X = 6.8; female range: 5.3 - 7.3, X = 6.6. Width at connexiva (mm) – Range: 4.9 - 7.5, X = 6.1; male range: 4.9 - 6.6, X = 5.8; 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 surrounding abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob 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. Measurements – Males: n = 4; females: n = 9.

54 Length (mm) – Range: 15.9 - 19.0, X=17.7; male range: 15.9 - 19.0, X=17.7; female range: 16.2 - 18.9, X = 17.7. Width at pronotum (mm) – Range: 5.8 - 7.9, X = 6.9; male range: 6.0 - 7.6, X = 6.9; female range: 5.8 - 7.9, X = 6.9. Width at connexiva (mm) – Range: 4.9 - 6.8, X = 6.1; male range: 4.9 - 6.1, X = 5.6; 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. Measurements – Males: n = 2; females: n = 4. 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 darkbrown 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 abdominal segment IV spiracle not raised as trigonal area. Metacoxal knob lacking. Proand 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. Measurements – Males: n = 10; females: n = 10. Length (mm) – Range: 19.7 - 23.3, X = 21.3; male range: 19.7 - 23.3, X = 21.6; female range: 20.3 - 22.2, X = 21.0. Width at pronotum (mm) – Range: 6.6 - 8.1, X = 7.3; male range: 6.6 - 7.7, X = 7.2; female range: 6.7 - 8.1, X = 7.3. Width at connexiva (mm) – Range: 6.5 - 8.8, X = 7.5; male range: 6.5 - 8.1, X = 7.2; 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 doublepoint. Anterior face of metatibiae concolorous with posterior face; metatibiae slightly arcuate; external apical tubercle present in males. Measurements – Males: n = 10; females: n = 10. Length (mm) – Range: 15.5 - 18.8, X = 17.1; male range: 15.6 - 18.8, X = 17.3; female range: 15.5 - 17.6, X = 16.9. Width at pronotum (mm) – Range: 5.0 - 6.4, X = 5.7; male range: 5.0 - 6.4, X = 5.6; female range: 5.3 - 6.2, X = 5.9. Width at connexiva (mm) – Range: 4.6 - 6.5, X = 5.3; male range: 4.6 - 5.3, X = 5.0; 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. Measurements – Males: n = 10; females: n = 10.

63 Length (mm) – Range: 15.4 - 19.6, X = 17.5; male range: 15.4 - 18.5, X = 17.1; female range: 17.0 - 19.6, X = 18.0. Width at pronotum (mm) – Range: 5.9 - 7.5, X = 6.6; male range: 5.9 - 7.2, X = 6.5; female range: 6.2 - 7.5, X = 6.7. Width at connexiva (mm) – Range: 7.9 - 9.6, X = 8.7; male range: 7.9 - 9.4, X = 8.5; 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. Measurements – Males: n = 10; females: n = 10. Length (mm) – Range: 19.0 - 23.8, X = 20.7; male range: 19.2 - 23.8, X = 20.7; female range: 19.0 - 22.6, X = 20.7. Width at pronotum (mm) – Range: 5.7 - 7.6, X = 6.8; male range: 5.7 - 7.5, X = 6.7; female range: 6.4 - 7.6, X = 6.9. Width at connexiva (mm) – Range: 5.1 - 7.1, X = 6.3; male range: 5.1 - 6.6, X = 5.9; 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. Laterally projecting jugal shelf present beneath base of antennal segment I. Beak 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. Length (mm) – Range: 16.4 - 20.9, X = 18.9; male range: 16.4 - 20.9, X = 18.7; female 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. Width at connexiva (mm) – Range: 7.5 - 9.8, X = 8.9; male range: 7.5 - 9.7, X = 8.8; 
      

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 References

Adler, P. H. and A. G. Wheeler Jr. 1984. Extra-phytophagous food sources of Hemiptera:Heteroptera: bird droppings, dung, and carrion. Journal of the Kansas Entomological Society. 57(1): 21-27. Amyot, C. J. B. and A. Serville. 1843. Histoire naturelle des insectes. Hémiptères. Librarie Encyclopédique de Roret. Paris: Fain et Thunot. Baranowski, R. M. and J. A. Slater. 1986. Arthropods of Florida and neighboring land areas volume 12: Coreidae of Florida (Hemiptera: Heteroptera). Gainesville: Florida Department of Agriculture and Consumer Services. 88 pp. Barber, H. G. 1914. Insects of Florida. II. Hemiptera. Bulletin of the American Museum of Natural History. 33: 495-535. Blatchley, W.S. 1926. Heteroptera or true bugs of eastern North America with special reference to the faunas of Indiana and Florida. Indianapolis: The Nature Publishing Company. 1116 pp. Blöte, H. C. 1938. Catalogue of the Coreidae in the Rijksmuseum van Natuurlijke Historie, part IV: Coreinae, third part. Zoologische Mededeelingen 20: 275-308. Brailovsky, H. 1993. Género nuevo y especies nuevas de Coreidos neotropicales (Hemiptera-Heteroptera-Coreidae: Acanthocerini, Chariesterini, Coreini, Discogasterini, Leptoscelidini y Nematopodini). Anales de Instituto Biológica, Universidad Nacional Autónomá México. Ser. Zoología. 64(2): 109-137. Brailovsky, H. and E. Barrera. 1983. Descriptión de dos nuevas especies Sudamericanas de la tribu Nematopodini (Hemiptera-Heteroptera-Coreidae). Anales de Instituto Biológica, Universidad Nacional Autónomá México. Ser. Zoología. 54(1): 69-77. Brailovsky, H. and E. Barrera. 1984. Hemíptera-Heteróptera de México XXXIII. El género Piezogaster Amyot y Serville con descriptión de dos nuevas especies (Coreidae-Nematopodini). Anales de Instituto Biológica, Universidad Nacional Autónomá México. Ser. Zoología. 55(1): 133-154. Brailovsky, H. and E. Barrera. 2000. Four new species of Neotropical Coreidae (Insecta: Hemiptera: Heteroptera). Reichenbachia. Staatliches Museum für Tierkunde Dresden. 33(34): 271-280. Burmeister, H. C. C. 1835. Handbuch der Entomologie, Zweiter Band. Besondere Entomologie. Ordnung Rhynchota. Abtheil 1: i-xii. Berlin: T. Enslin. 400 pp.

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Dallas, W. S. 1852. List of the specimens of Hemipterous insects in the collection of the British Museum. Part II. London: Taylor & Francis Inc. pp. 369-592. Dallwitz, M. J., T. A. Paine, and E. J. Zurcher. DELTA Primer1980-1999. DELTA: Description Language for Taxonomy, v.1.01 (software). http://biodiversity.uno.edu/delta/. Distant, W. L. 1880-1893. Insecta. Rhynchota. Hemiptera-Heteroptera. Vol. 1. Biologia Centrali-Americana. London. 462 pp. [1881: 89-168; 1893: 329-462]. Distant, W. L. 1901. Rhynchotal notes IX. Heteroptera: Fam. Coreidae. Annals and Magazine of Natural History 7(7): 416-432. Environmental Systems Research Institute. 1992-1998. ArcView v. 3.1 (software). Environmental Systems Research Institute. Redlands, California. http://www.esri.com. Fabricius, J. C. 1803. Systema Rhyngotorum secundum ordines, genera, species, adjectis, synonymis, locis observationibus, descriptionibus. Brunsvigae: Apud Carolum Riechard vi. 314 pp. Henry, T. J. and R. C. Froeschner. 1988. Catalog of the Heteroptera, or true bugs, of Canada and the continental United States. New York: E.J. Brill. 958 pp. Herrich-Schäffer, G. H. W. 1842. Die wanzenartigen insekten. Nürnberg: C. H. Zeh'sche Buchhandlung. 6: 1-118. Laporte de Castelnau, F. L. N. 1833. Essai d'une classification systématique de l'ordre des Hémiptères (Hémiptères Hétéroptères Latr.). Magasin de Zoologie (1832). Lethierry, L. and G. Severin. 1894. Catalogue général des Hémiptères. Vol II. Hétéroptères: Coreidae, Berytidae, Lygaeidae, Pyrrhocoidae. Brussells: Musée Royal D'Histoire Natuelle de Belique. Montandon, A. L. 1897. Hémiptères-Hétéroptères de l'Equateur. Trois espèces nouvelles de la fam. Corediae. Bulletin de la Société des Sciences. Bucarest 3: 246-251. Montandon, A. L. 1899. Notes et description de trois nouvelles espèces Américaines. Bulletin de la Société des Sciences. Bucarest 8: 190-195. O'Shea, R. 1974. Two new combinations and a new synonymy in the Coreidae (Hemiptera-Heteroptera). Entomologist's Monthly Magazine. 109: 125.

74 O'Shea, R. 1980. A generic revision of the Nematopodini (Heteroptera: Coreidae: Coreinae). Studies on Neotropical Fauna and Environment 15: 197-227. O'Shea, R. and C. W. Schaefer. 1978. The Mictini are not monophyletic (Hempitera: Coreidae: Coreinae). Annals of the Entomological Society of America. 71: 776-784. Packauskas, R. J. 1994. Key to the subfamilies and tribes of the New World Coreidae (Hemiptera), with a checklist of published keys to genera and species. Proceedings of the Entomological Society of Washington. 96(1): 283-287. Say, T. 1825. American Entomology. I. Volume 2. Philadelphia. 112 pp. Schaefer, C. W. and P. L. Mitchell. 1983. Food plants of the Coreoidea (Hemiptera: Heteroptera). Annals of the Entomological Society of America. 76(4): 591-60?. Schaefer, C. W. and R. O'Shea. 1979. Host plants of three Coreine tribes (Hemiptera: Heteroptera: Coreidae). Annals of the Entomological Society of America. 72(4): 519-523. Stål, C. 1862. Hemiptera Mexicana enumeravit species-que novas descripsit. Stettin Entomologische Zeitung. 23(4-6): 273-281; 23(7-9): 289-325. Stål, C. 1867. Bidrag till Hemipterernas systematik. Öfversigt af Kongl. VetenskapsAkademiens Förhandlinger. 24(7): 491-560. Stål, C. 1870. Enumeratio Hemipterorum: Bidrag till en företeckning öfver alla hittills kända Hemiptera, jemte systematiska meddelanden. Part 1. Kongliga Svenska. Vetenskaps-Akademiens Handlingar 9(1): 1-232. Torre-Bueno, J. R. de la. 1941. A synopsis of the Hemiptera-Heteroptera of America north of Mexico. Part II. Entomologica Americana. 21(2): 41-122. Uhler, P. R. 1876. List of the Hemiptera of the region west of the Mississippi River, including those collected during the Hayden explorations of 1873. Bulletin of the U.S. Geologic and Geographic Survey of the Terrain (2nd Ser.). 5: 296-297. Van Duzee, E. P. 1909. Observations on some Hemiptera taken in Florida in the spring of 1908. Bulletin of the Buffalo Society of Natural Science. 9: 149-230. Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico, excepting the Aphididae, Coccidae and Aleurodidae. Berkeley: University of California Press. 902 pp. Walker, F. 1871. Catalogue of the specimens of Hemiptera Heteroptera in the collection 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. Transactions of the Wisconsin Acadamey of Sciences, Arts and Letters. 57: 163-188. Yonke, T. R. and J. T. Medler. 1969b. Description of immature stages of Coreidae. 3. Archimerus alternatus. Annals of the Entomological Society of America. 62(3): 477-480.

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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 136139: 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.