AN ABSTRACT OF THE THESIS OF

Kathryn Ann Phillips for the degree of Doctor of PhilosoPhY in Entomology presented on December 22, 1982

Title: A Taxonomic Revision of the Nearctic Soecies of

Apateticus Dallas and Herrich-Schaeffer

(: : ) Redacted for privacy Abstract approved: John D. Lattin

The Nearctic of the pentatomid genera

Apateticus Dallas and Podisus Herrich-Schaeffer were revised taxonomically. Approximately 8500 Nearctic specimens plus 500 extralimital examples were examined.

Five species of Apateticus and 9 of Podisus, including 1 incidental Nearctic species, are redescribed. Important characters for species discrimination include shape of the male claspers, length of the ventral abdominal spine, and characteristics of the pronotum. Female genitalic characters are not reliable for species-level distinctions within each . Re-evaluation of previously used characters and additional male and female genitalic traits support the idea that these genera should remain separate.

A cladistic analysis was performed on the species of each

genus.

One new species, Podisus brevispinus, is described.

The latter, heretofore known as P. modestus (Dallas), has

long been associated with a type specimen which is a

synonym of P. maculiventris (Say). Examination of

Rhaphigaster aggressor Walker, listed as synonymous with

P modestus, showed the former to be a Neotropical Podisus.

The following synonymies are suggested (junior

synonym first): A. crocatus (Uhler) = A. bracteatus (Fitch),

P. pallens (Sega) = P. maculiventris (Say) (reaffirmed),

P. modestus (Dallas) = P. maculiventris (Say), and P. rubromaculatus Distant = P. acutissimus Stk.

Podisus maculiventris, P. brevispinus, P. placidus

Uhler, P. serieventris Uhler, and hEattLizas bracteatus are widely distributed in the and Canada.

A. cynicus (Say) occurs from northeastern

south to Florida and west to Arizona. P. acutissimus, P.

sagitta (F.), and A. lineolatus (H.-S.) occur in the southern U. S., Mexico, Central America, and the Caribbean.

P. mucronatus Uhler is confined to Florida and Cuba, while

P. congrex (Stil) occurs from central Mexico into South

America. Localized distributions are found in A. anatarius

Van Duzee, A. marginiyentris (Stg.1), and P. fretus, Olsen. Apateticus and Podisus prey on a wide variety of , notably larval , , and

Coleoptera. Podisus maculiventris is being investigated as a biological control agent of lepidopterous pests of in the United States and has been introduced into

Europe and Asia with varying success. Lists of known prey are presented and major biological literature for each species is reviewed. A Taxonomic Revision of the Nearctic Species of Apateticus Dallas and Podisus Herrich-Schaeffer (Heteroptera: Pentatomidae: Asopinae)

by

Kathryn Ann Phillips

A THESIS

submitted t

Oregon State University

in partial fulfillment of the requirements for the degree of

Doctor of Philosophy

Commencement June 1983 APPROVED:

Redacted for privacy

Professor/6f Entomology in charge ofmajor

Redacted for privacy Head of Department of Entomolo

Redacted for privacy Dean of Gradua scnool ci

Date thesis is presented_ December 22, 1982

Typed by Kathryn Ann Phillins ACKNOWLEDGEMENTS

I express my sincere appreciation to the many people who assisted and encouraged me throughout this undertaking.

Financial support was provided through a Resident

Instruction graduate teaching assistantship from September,

1979 through December 1980 and by the Science Technical Advisory Fund through a research assistantship from

January 1981 through December 1982. I received support for computer services for unsponsored research from the

Oregon State University Computer Center.

Type specimens were lent by the American Museum of Natural History, New York, (AMNH), Dr. R. T. Schuh; British Museum (Natural History), London (BM), Dr. W. R. Dolling;

California Academy of Sciences, San Francisco, (CAS), Dr.

P. H. Arnaud, Jr.; Museum of Comparative Zoology, Harvard

University, Cambridge, Massachusetts, (MCZ), Mr. Alfred F.

Newton, Jr.; Naturhistorisches Museum, Vienna, Dr. A.

Kaltenbach; Naturhistoriska Riksmuseet, Stockholm, Dr. P.

Lindskog; United States National Museum, Washington, D. C.,

(USNM), Mr. T. J. Henry.

The following institutions and individuals kindly made loans of specimens: American Museum of Natural

History, New York (AMNH), Dr. R. T. Schuh; Arizona State University, Tempe (ASU), Dr. F. F. Hasbrouck; California

Academy of Sciences, San Francisco, (CAS), Dr. P. H.

Arnaud, Jr.; Carnegie Museum of Natural History,

Pittsburgh, PA, (CMNH), Dr. G. Ekis; State

University, Fort Collins, (CSU), Dr. W. D. Fronk; Cornell

University, Ithaca, NY, (CU), Dr. L. L. Pechuman; Joe E.

Eaer Collection, Tampa, FL, (JE), Dr. J. E. Eger; Illinois

Natural History Survey, Urbana, (INH), Mr. D. Voegtlin;

Illinois State Museum, Springfield, (ISM), Dr. E. D.

Cashatt; Iowa State University, Ames, (ISU), Dr. R. E.

Lewis; Louisiana State University, Baton Rouge, (LSU),

Dr. J. B. Chapin; State University, East Lansing,

(MIS), Dr. R. L. Fischer; Montana State University,

Bozeman, (MSU), Ms. S. D. Rose; Museum of Comparative Zoology, Harvard University, Cambridge, MA, (MCZ), Ms.

M. Hathaway; Natural History Museum of Los Angeles County,

CA, (LACM), Dr. C. L. Hogue; North Carolina State

University, Raleigh, (NCSU), Ms. C. Parron; North Dakota State University, Fargo, (NDSU), Dr. E. U. Balsbaugh, Jr.;

Northern Arizona University, Flagstaff, (NAU), Dr. C. D.

Johnson; Ohio State University, Columbus, (OHS), Dr. C. A.

Triplehorn; Oklahoma State University, Stillwater, (OKS),

Dr. W. A. Drew; Oregon State University, Corvallis, (OSU),

Dr. J. D. Lattin; Peabody Museum, Yale University, New

Haven, CT, (YLU), Mr. D. G. Furth; Pennsylvania State

University, University Park, (PSU), Dr. K. C. Kim; Purdue University, West Lafayette, IN, (PDU), Mr. A. V. Provonsha;

San Diego Society of Natural History Museum, CA, (SDCM),

Mr. D. K. Faulkner; Gary M. Stonedahl Collection,

Corvallis, OR, (GMS), Dr. G. M. Stonedahi; Texas A & M

University, College Station, (TAM), Dr. J. C. Schaffner;

Texas Tech University, Lubbock, (TTU), Dr. O. F. Francke;

Donald B. Thomas, Jr. Collection, Lincoln, NE, (DBT), Dr.

D. B. Thomas, Jr.; University of Alberta, Edmonton, (UAL),

Dr. G. E. Ball, Mr. D. Shpeley; University of Arizona,

Tucson, (UAZ), Dr. F. G. Werner; University of British

Columbia, Vancouver, (UEC), Mr. S. G. Cannings; University of California, Berkeley, (UCB), Dr. J. A. Powell, Mr. J.

B. Whitfield; University of California, Riverside, (UCR),

Mr. S. I. Frontier; University of Connecticut, Storrs,

(UCT), Ms. J. E. O'Donnell; University of Georgia, Athens,

(UGA), Dr. C. L. Smith; University of Idaho, Moscow, (UID),

Dr. W. F. Barr; University of Kansas, Lawrence, (KU), Dr.

P. D. Ashlock; University of Kentucky, Lexington, (UKY),

Dr. P. H. Freytag; University of Michigan, Ann Arbor,

(UMI), Dr. T. E. Moore, Mr. M. F. O'Brien; University of

Minnesota, St. Paul, (UNN), Dr. P. E. Clausen; University of Missouri, Columbia, (UMO), Mr. E. G. Riley; University of Nebraska, Lincoln, (UNL), Dr. B. C. Ratcliffe; University of Wisconsin, Madison, (UWI), Mr. S. Krauth; University of

Wyoming, Laramie, (UWY), Dr. R. J. Lavigne; Virginia

Polytechnic Institute and State University, Blacksburg, (VPI), Dr. M. Kosztarab; Washington State University,

Pullman, (WSU), Dr. R. S. Zack.

I also thank Dr. Kenneth V. Yeargan, University of

Kentucky, Lexington, for providing live eggs and preserved nymphs of Podisus maculiventris. Dr. Donald B. Thomas, Jr. generously allowed me the use of his notes on Podisus, and offered useful suggestions. Dr. Joe E. Eger also gave

advice and suggestions, and supplied slides of Apateticus.

To my colleagues, Dr. Gary M. Stonedahi and Mr. Jerry Cassis I am grateful for ideas, many discussions, and help. Jerry also prepared the scanning electron micrograph

of Anateticus bracteatus shown in Figure 16.

To the members of my thesis committee, Drs. Arnold P.

Appleby, Kenton L. Chambers, Paul L. Farber, Jeffrey C.

Miller, and Paul W. Oman, I express ray sincere thanks. My major professor, Dr. John D. Lattin, suggested this project

and allowed me full use of his excellent library. He also

provided encouragement, advice, and friendship throughout

this project's duration. To him I owe special gratitude.

Special appreciation goes to Mr. Michael E. Szmania

for encouragement and moral support over the last 3 years.

To my parents, Richard B. and Frances K. Phillips, who

throughout my life have taught me the value of knowledge

and provided me overy opportunity to pursue it, I dedicate

this thesis. TABLE OF CONTENTS

INTRODUCTION

LITERATURE REVIEW 6

MATERIALS AND METHODS 9

BIOLOGY AND ECONOMIC IMPORTANCE 12

DISTRIBUTION 17

METHOD OF CLASSIFICATION 20 DISCUSSION OF TAXONOMIC CHARACTERS 23

TAXONOMY 31 Genus Apateticus Dallas 34

Key to the Species of Apateticus 36

Subgenus Apateticus Dallas 38 Apateticus lineolatus (Herrich - Schaeffer) 38

Apateticus marainiventris (Stga) 43

Subgenus Apoecilus St&l. 46

Apateticus bracteatus (Fitch) 47 Apateticus anatarius Van Duzee 56

Apateticus cynicus (Say) 9

Genus Podisus Herrich - Schaeffer 67

Key to the Nearctic Species of Podisus 70

Subgenus Tylospilus St1.1 72

Podisus acutissimus Stcal 72 Subgenus Podisus Herrich - Schaeffer 82

Podisus mucronatus Uhler 83

Podisus maculiventri (Say) 87 Podisus serieventris Uhier 114

Podisus concfrex (St&l) 122

Podisus brevispinus, new species 125

Podisus fretus Olsen 138 Podisus placidus Uhier 142

Podisus sagitta (Fabricius) 152

CONCLUSIONS 159

LITERATURE CITED 226

APPENDICES 244 LIST OF FIGURES

Fiaure Page,

1 Apateticus lineolatus (Herrich - Schaeffer) 209

2 Apateticus marginiventris (St&l) 209

3 Apateticus bracteatus (Fitch) 211

4 Apateticus anatarius Van Duzee 211

5 Apateticus cynicus (Say) 213

6 Podisus mucronatus Uhler 213

7 Podisus acutissimus StL1 215

8 Podisus maculiventris (Say) 215

9 Podisus serieventris Uhler 217

10 Podisus congrex (Staff) 217

11 Podisus brevispinus, new species 219

12 Podisus fretus Olsen 219

13 Podisus placidus Uhler 221

14 Podisus saaitta (Fabricius) 221

15 a,b: Rostrum, Supputius cincticeps (StL.), P. maculiventris.

cod: Head, A. lineolatus, P. maculiventris.

e,f: Conjunctival appendages, A. bracteatus, A. lineolatus.

g,h: Median penial lobes, A. bracteatus, P. maculiventris.

i,j: Right male genital plate, P. acutissimus, P. maculiventris.

k,1: Duct of metathoracic scent gland, P. acutissimus, P. maculiventris. 223 Fiaure page,

16 Scanning electron micrograph of abdomen of Apateticus bracteatus, male, showing setose, rugulose areas on sternites IV - VI. 40 X. 225 LIST OF TABLES

Table Page

1 Summary of life history data for Apateticus and Podisus expressed in days. 162

2 Prey records for Apateticus and Podisus. 165

3 Parasitoids attacking species of Podisus. 177 LIST OF MAPS

Page

1 Distribution of Apateticus lineolatus (Herrick - Schaeffer) 181

2 Distribution of Apateticus marginiventris (StW 183

3 Distribution of Apateticus bracteatus (Fitch) 185

4 Distribution of Apateticus anatarius Van Duzee 187

5 Distribution of Apateticus cvnicus (Say) 189

6 Distribution of Podisus mucronatus Uhler 191

7 Distribution of Podisus acutissimus St&l 193

8 Distribution of Podisus maculiventris (Say) 195

9 Distribution of Podisus serieventris Uhler 197

10 Distribution of Podisus conarex (St11) 199

11 Distribution of Podisus brevispinus, new species 201

12 Distribution of Podisus fretus Olsen 203

13 Distribution of Podisus placidus Uhler 205

14 Distribution of (Fabricius) 207 A TAXONOMIC REVISION OF THE NEARCTIC SPECIES OF APATETICUS DALLAS AND PODISUS HERRICH-SCHAEFFER (HETEROPTERA: PENTATOMIDAE: ASOPINAE)

INTRODUCTION

The family Pentatomidae, commonly known as stink bugs, is one of the largest in the order Heteroptera. Depending on the authority, this cosmopolitan group is divided into a number of subfamilies. I have followed the classification of Torre-Bueno (1939). There are approximately 57 genera in America north of Mexico, the majority in the subfamily Pentatominae, the members of which are phytophagous. Some pentatomine species, such as

Nezara viridula (L.), the southern green stink bug, and species of Euschistus are of economic importance (Mundinger and Chapman 1932; Borden and Madsen 1951; Rings 1957; Todd and Herzog 1980). Members of the subfamily Asopinae, to which Apateticus Dallas and Podisus Herrich - Schaeffer belong, are predaceous.

Pentatomids exhibit great diversity in size and coloration. Among the smallest North. American representatives are Neottialossa Kirby, approximately 5MM long, whereas individuals longer than 20 mm are found in 2

Apateticus and Alcaeorrhynchus Bergroth. Browns and greens are predominant hues among our fauna, but striking red and black or blue coloration is typical of suchgenera as Murciantia StalOplomus Spinola, Perillus Stall

Stiretrus Laporte, and Zicrona Amyot & Serville. All pentatomids have well-developed scent glands that produce pungent, volatile secretions, some of which have been chemically analyzed (Blum et al. 1960; Blum 1960;

Waterhouse et al. 1961; Remold 1963; Ishiwatari 1974, 1976; Aldrich et al. 1978; Staddon 1979).

The Nearctic Pentatomidae are fairly well known. Basic life history studies and descriptions of nymphs and eggs have been published for many species (Esselbaugh 1946,

1948; DeCoursey and Esselbaugh 1962; McPherson 1982, references therein). However, many more remain to be studied. This family has attracted a host of workers, whether because of economic importance, interest in chemical composition of scents, or unabashed curiosity.

My experience has been that pentatomids are easy to rear and interesting to observe.

The subfamily Asopinae is of interest to workers in pest management. Torre-Bueno (1939) lists 14 genera for America north of Mexico. Apateticus and especially Podisus are among the most common. One accidentally introduced European species, bidens

(L.), has been reported preyingon (Hartig), 3 the introduced pine , on pine in Vermont (Cooper

1967), Pyrrharctia isabella (J. E. Smith) in New Hampshire

(Lattin and Donahue 1969), and Gilpinia fruteorum (F.), an introduced sawfly, in Quebec (Kelton 1972). Currently,

Podisus maculiventris (Say) is under evaluation as a biological control agent in conjunction with other predaceous Heteroptera (Irwin and Shepard 1980).

Species of Apateticus and Podisus, like all pentatomids, pass through an egg stage and 5 nymphal instars prior to becoming adults. First instar Asopinae imbibe liquids from plants or unhatched eggs, but are not predaceous (Plummer and Landis 1932; Prebble 1933; Coppel and Jones 1962; LeRoux 1965; Oetting and Yonke 1971;

Tostowaryk 1971; Stoner et al. 1974a,b; pers. obs.).

Instars II - V and adults are predators of soft-bodied lepidopterous and coleopterous larvae.Among these prey are serious agricultural and silvicultural pests such as

Epilachna varivestis Mulsant, the Mexican bean

(Howard and Landis 1936; Waddill and Shepard 1975a),

Leptinotarsa decemlineata (Say), the (Warren and Wallis 1971), Heliothis zea (Boddie), the corn earworm (Lopez et al. 1976), Lymantria dispar

(L.), the gypsy (Kirkland 1898), and Diprion similis

(Hartig), the introduced pine sawfly (Coppel and Jones

1962). Members of Apateticus overwinter as eggs (Whitmarsh

1916; Downes 1920; Prebble 1933; Stone 1939; Jones and 4

Coppel 1963). Adults of Podisus spp. are the overwintering

stage (Warren and Wallis 1971).

My interest in pentatomids began while doing master's research on 2 species of Euschistus suspected of injuring apples in Georgia. Extensive collecting in orchards and

surrounding fields, rearing of nymphs, and identifying

adults made me aware of taxonomic difficulties with some genera. At the beginning of my tenure at Oregon State

University, Dr. J. D. Lattin, my major professor, wisely advised that I concentrate my taxonomic interests in the

Asopinae and suggested an examination of Apateticus and

Podisus.

There are over 65 species in Apateticus and Podisus

combined. I decided to revise these genera in the

Nearctic region addressing the following:

1. Assessing morphological evidence for maintaining or

reuniting the genera.

2. The status of A. crocatus (Uhler), recognized as a

separate species by some, but treated as a variety of

A. bracteatus (Fitch) by Kirkaldy (1909).

3. The distribution of P. maculiventris (Say) and status of P. pallens (Stal), included by Van Duzee (1917) as

a synonym of the former.

4. Clarification of the confusion between P. maculiventris, and P. serieventris (see Prebble 1933, p. 2). 5

5. Clarification of confusion between P. saqitta (F.) and P. fuscescens (Dallas).

6. Clarification of species distributions.

7. Evaluation of male and female genitalic characters in

classification.

8. A cladistic analysis of the species in each genus.

9. Compilation of biological literature for each species,

facilitating assessment of research needs by workers

in biological control and integrated pest management. 6

LITERATURE REVIEW

Many general treatments of the Pentatomidae are available because of the group's interest to researchers.

Uhler (1886), Gillette and Baker (1895), Summers (1898),

Van Duzee (1904), Zimmer (1911), Parshley (1915), Hart

(1919), Blatchley (1926), Downes (1926), and Baker (1927a) provide keys and descriptions of all or part of the North

American species. Torre-Bueno (1939) published a key to all North American species including host data and a glossary. Stoner (1916, 1917, 1925) gives species lists and distributional data for Iowa and the Douglas Lake area in Michigan. Froeschner (1941), McPherson (1970, 1982),

Hoffman (1971), and Furth (1974) continue in this tradition but restrict their treatments to Missouri, Michigan, the northeastern U. S., Virginia, and Ohio, respectively. All of these keys focus mainly on external characters for species discrimination.

McPherson's (1982) book on the of the northeastern U. S. summarizes life history data and distribution for each species and offers a good introduction to the literature on this group. Slater and Baranowski's

(1978) illustrated key includes all the families but not all of the genera in the order (Heteroptera) of 7

North America.

Studies evaluating the value of genitalia and other characters in the classification of the Pentatomidae and the Heteroptera in general also exist. Baker (1931) examined male genitalia in Canadian Pentatomidae, including

A. bracteatus, P. maculiventris, P. brevispinus (as P. modestus (Dallas)), and P. placidus Uhler. Each structure was described and considered in the light of evolutionary development within the family. McDonald (1966), in a study of the genitalia of North American Pentatomoidea, examined and illustrated the males of 85 species and females of 80 species, including A. bracteatus, A. lineolatus (H.-S.),

P. acutissimus St&l, and P. maculiventris. Included is a discussion of interrelationships within the superfamily

Pentatomoidea. Schaefer (1968) assessed homologies among female pentatomoid reproductive structures.

General works examining the value of genitalia in classification of the Heteroptera include those of Singh-

Pruthi (1925), Dupuis (1955), Pendergrast (1957), and

Scudder (1959). Dupuis (1955) and Pendergrast (1957) considered both sexes, Singh-Pruthi (1925) examined males, and Scudder (1959) evaluated female reproductive structures.

Other studies considering evolutionary trends and classification in Heteroptera include Southwood (1956) on egg structure, Cobben (1968, 1978) on eggs, development, mouthparts, and feeding, Leston (1958) on chromosome number 8 in relation to the systematics of the Pentatomorpha, and

Leston et al. (1954) on the classification of terrestrial Heteroptera. 9

MATERIALS AND METHODS

Just over 8500 specimens of ADateticus and Podisus

were examined for this study. Additionally, 500 examples of extralimital species and about 50 from related asopine

genera were seen. Many institutions lent specimens; a

list of institutions, curators, and label codes is

presented in the acknowledgements. Types were kindly

provided by several institutions whichare also listed in

the acknowledgements. I made collections by beating and sweeping in Oregon, Georgia, and Colorado.

Upon receipt of a loan, a label bearing a 2 - 4

letter code representing the lending institution was pinned below each insect. Initially the insects were sorted by

state or province. Once the arrival of specimens slowed, those in each state were divided into "species" based on external morphology. Next, each "species" was sorted by county withineach state so that plotting of distributions would be facilitated, Label data for each specimen were recorded as follows: COUNTRY: State: County: Locality (institution). Types were kept separate from other specimens.

638 males and 541 females, including extralimital species and related genera were dissected. Examples of 10 each species were selected from all parts of its range, beginning with the northwest and progressing eastward.

Candidates for dissection were tagged with a numbered label attached to the pin. Females and males were numbered separately. Each specimen was soaked in lukewarm water containing a small amount of soap and rinsed in water.

With jeweler's forceps I removed the male genital cup or the entire abdomen of the female. Each cup or abdomen was placed in a small glass tube containing 10% KOH, inserted into a water bath, and boiled for 7 - 10 minutes depending on the specimen's size. After boiling, parts were removed from the KOH, dipped briefly in dilute acetic acid, and placed separately in depression plates containing distilled water.

Dissection was done under 27X and 54X magnification using a Spencer dissecting microscope equipped with a calibrated ocular grid. For males, shape of the genital cup was noted. The proctiaer and claspers were then examined for differences in shape and orientation. The aedeagus, if not inflated, was pulled and pushed out from behind the proctiger.Aedeagal differences were readily apparent between genera or subgenera but not sufficiently distinct among species to be used in their discrimination. Sketches of the right clasper, genital plates, and aedeagus were made for each specimen and recorded along with locality data and measurements (see "Discussion of Taxonomic 11

Characters," p. 23).

For females, width and length of the first gonocoxa were measured. Next the dorsal and ventral halves of the abdomen were teased apart between the connexiva and terga

Any extraneous material not dissolved by boiling was gently pulled away from the spermatheca. Sketches of the spermatheca, bulb, and gonocoxae were made and the sclerotized rod inside the spermathecal dilation was measured. Measurement and locality data for each specimen were recorded alongside.

Each dissection was placed in a microvial containing glycerin and pinned below the corresponding insect. Any tachinid larvae or unlaid eggs found with a female's abdomen were placed with the dissection in the microvial.

Microvials received numbers matching those on the numbered paper tags.

Dissected specimens were also measured externally. In species where fewer than 100 individuals were examined all were measured, regardless of dissection. Every type specimen was also measured. All data are given in milli- meters and measurements are described in the section on taxonomic characters (p. 23).

Drawings were made using the same microscope at 18X,

27X, or 54X magnification and are to scale. Structures were aligned with the ocular grid and drawn on prepared tracing paper clipped to a large grid. 12

BIOLOGY AND ECONOMIC IMPORTANCE

Esselbaugh (1946) studied and described the eggs of various pentatomid species. The eggs are laid in masses and are characteristically barrel-shaped with a row of slender chorionic processes encircling the top. At hatching, movements of the nymph exert pressure on the egg burster which causes the chorion to break along a circular suture just interior to the chorionic processes. This lid, or operculum, is flipped up as the nymph crawls out.

Nymphs pass through 5 successively larger instars prior to becoming adults. First instars usually remain clustered on or near the hatched eggs until after the first molt. Immature pentatomids have 3 pairs of dorsal abdominal scent glands that open between tergites III-IV,

IV-V, and V-VI (DeCoursey and Esselbaugh 1962). The glands secrete a liquid noted for its odor and responsible for the common name of these insects, stink bugs. In

Nezara viridula (L.) nymphs, the scent substance acts as either an alarm or aggregation pheromone, depending on the concentration (Ishiwatari 1974, 1976).

Adult pentatomids possess metathoracic and dorsal abdominal scent glands opening between tergites III-IV

(Aldrich et al. 1978). The secretions of the glands of 13 some species have been analyzed chemically and found to contain a diversity of substances (see Staddon 1979 fora review).

Pentatomids are attacked by hymenopteran egg parasitoids and parasitized as late stage nymphs and adults by of the family . Table 3 shows the parasitoids known to attack several species of Podisus.

Parasitized eggs turn black and each emerges after cutting a circular opening in the top of the stink bug egg. Tachinid flies lay small, pearly white eggs externally on the bug's body. The larva uses its mouth hooks in a rasping motion to bore directly into the pentatomid's body (Worthley 1924). The larva develops inside its host and emerges, usually through the bug's genital or excretory orifice, to pupate in the soil. The stink bug dies shortly thereafter.

The predaceous Asopinae can be separated from the phytophagous pentatomids by the presence of bucculae that do not extend along the length of the first rostral segment. Because of their predatory nature, there is growing interest in using species of Asopinae in integrated pest management programs. For example,

Stiretrus anchorago (F.) is being evaluated as an agent in the control of soybean insects (Waddill and Shepard

1975a; Irwin and Shepard 1980). Perillus bioculatus (F.) has been introduced into various parts of Europe and Asia 14 to control the Colorado potato beetle (Leptinotarsa decemlineata (Say)) with varyingsuccess (Briand 1936;

Tremblay 1966; Lipa 1976; Jermy 1980). P. bioculatus has been the subject of many laboratory studies (Shagov 1968, 1976, 1977a,b; Jasic 1970, 1975).

Apateticus spp. and Podisus spp. are polyphagous predators in a diversity of habitats, usually found in small numbers. Table 2 lists prey of these insects.

These pentatomids pass through 5 nymphal instars, the first of which subsists on plant juices, water, or unhatched eggs (Downes 1920; Plummer and Landis 1932;

Prebble 1933; Coppel and Jones 1962; LeRoux 1965; Oetting and Yonke 1971; Tostowaryk 1971; pers. obs.). P. maculiventris has been introduced into Europe, Korea, and Japan to control the Colorado potato beetle and (Couturier 1938; Sazonova et al. 1976; Hokyo and

Kawauchi 1975; Kim et al. 1968). P. placidus was introduced into Yugoslavia for fall webworm control but did not overwinter successfully (Tadi6 1975).

P. maculiventris is being studied as a predator of soybean pests in the United States (Waddill and Shepard

1975a,b; Irwin and Shepard 1980).Although scarce and widely dispersed, this predator consumes larger prey than do other smaller, more abundant heteropteran predators such as Nabis sp., Geocoris sp., or Onus sp.

(Irwin and Shepard 1980). A South American species, 15

Podisus niqrispinus (Dallas), is also a predator in soybean agroecosystems (Irwin and Shepard 1980).

Podisus serieventris Uhler is generally found in coniferous forests where it preys on larvae of and forest Lepidoptera (see Table 2). Dahlsten et al.

(1977) reported that this species tended to respond to density of larvae of Orqyia pseudotsuqata (McDunnough), the Douglas-fir tussock moth, but more investigations are needed to confirm this. P. brevispinus preys on a wide variety of insects (see Table 2) and was a factor in controlling an outbreak of spp. on jack-pine in Quebec (Tostowaryk 1971).

Podisus saqitta (F.) and Apateticus lineolatus were the first and second most abundant pentatomid predators of the , Epilachna varivestis Mulsant, in Mexico (Plummer and Landis 1932). These predators were not well-synchronized with the prey, as their greatest abundance occurred after the bean beetle population peaked (Plummer and Landis 1932).

Evans (1982a) also noted a lack of synchrony of P. maculiventris with Trirhabda spp. (Chrysomelidae) in

Solidaqo sp. stands near Ithaca, NY. This author postulated that, as a generalist predator, P. maculiventris may be adapted to seeking outbreaks.

The role of other Apateticus spp. and Podisus spp. as predators is not known. Scattered prey records exist 16 (see Table 2) but expansion and re-examination of the roles of these generalist predators is critical to integrated pest management programs. Once this is done, ways to enhance the effectiveness of these insects may be developed. 17

DISTRIBUTION

The family Pentatomidae is of worldwide distribution.

Approximately 57 genera occur in America north of Mexico

(Torre-Bueno 1939). The predaceous subfamily Asopinae is also cosmopolitan and consists of over 50 genera and 300 species, most of which are New World (Kirkaldy 1909).

Gapud (1981) has completed a study in which he performed a cladistic analysis of the world Asopinae, but I have not yet seen a copy of the dissertation. Podisus, the largest genus (approximately 68 species) is confined to the New

World. Apateticus is among the smaller genera (5 species) and is also strictly New World.. The majority of Podisus species are Neotropical, while most species of Apateticus, are Nearctic.

Species distributions within Podisus and Apateticus overlap. A common pattern is a transcontinental distribution north of a line extending along the Ohio

River and west to the southern . The northern limits are southern or middle latitudes of Canada.

This distribution is seen, with variations, in Podisus brevispinus (Map 11), P. placidus (Map 13), P. serieventris

(Map 9), P. maculiventris (Map 8), and Apateticus bracteatus (Map 3). My estimates of Canadian distributions 18 are probably conservative, as the number of Canadian specimens examined was much smaller than that of U. S. specimens. The 5 aforementioned species vary in the extent of their distributions to the south. P. maculiventris, the most widespread species, occurs farther south than the other 4 (Map 8).

Habitat differences among P. brevispinus, P. placidus,

P. maculiventris, and P. serieventris do not appear sharply delineated. Coppel and Jones (1962) found all 4 species associated with an outbreak of Diprion similis

(Hartig) in Wisconsin. Evans (1979) found all but P. serieventris preying on the same colony of tent caterpil- lars. He observed that each species occupied a different position within or outside the tent.

Several species of Podisus and Apateticus, while occurring in the Nearctic region, have more extensive

Neotropical ranges. This distribution is generally from the southern U. S. throughout Mexico, Central America, and northern . P. fretus Olsen and A. anatarius

Van Duzee, 2 Nearctic species, have very limited distributions and are rarely collected.

Distributional studies should focus on a finer resolution of the habitat with which each species is associated. General habitat preferences such as agro- ecosystem or forest association, are known, but the degree to which a single population or even one individual is 19 confined to a particular habitat has been little studied.

This type of knowledge, coupled with known biogeographic history of North and South America, morphological, behavioral, and ecological data would generatea clearer understanding of the interrelationships among the species of Apateticus and Podisus. 20

METHOD OF CLASSIFICATION

Species were delineated on the basis of morphological characters as noted in the descriptions. Characters considered are described in the section on discussion of taxonomic characters (p. 23). A cladistic analysis was performed using the Wagner 78 Maximum Parsimony program of

Dr. James S. Farris, State University of New York,

Stonybrook, on the Oregon state University CDC Cyber computer. This program produces a tree of greatest parsimony for the fewest character changes.Relationships depicted therein are based solely on shared derived characters. Sister groups originating at each node share a hypothetical ancestor with characters apomorphic with respect to those of taxa to the left of that node on the tree. Also produced, depending on options selected, are a list of changes for each character, a table of ranges of character states, lengths, and consistency indices. If desired, a list of synapomorphies for each taxon and stem can also be generated, Discussions of and references to the Wagner method can be found in Schuh and Polhemus

(1980).

Mr. John Holt of the Oregon State University Computer

Center wrote a program to randomize columns and rows of 21 the data matrices before analysis by the Wagner 78 program.

This was used subsequent to each initial run of the latter.

Preliminary to the analysis was selection of an outgroup. I chose 10 genera of Asopinae with Nearctic and/ or Neotropical representatives (including Apateticus and

Podisus) and constructed a matrix of 14 bi- or multistate characters for analysis using the "mid" option of the

Wagner program. This causes the tree to be rooted between the 2 most divergent taxa. Through 10 runs of this matrix

4 genera consistently appeared together: Apateticus

Dallas, Podisus Herrich - Schaeffer, Supputius Distant, and Tynacantha Dallas. I analyzed the genera separately and obtained a tree rooted between Apateticus and

Tynacantha with Podisus and Supputius in between. Based on these results, Podisus maculiventris was arbitrarily selected as the outgroup for Apateticus and Supputius cincticeps (StL) was chosen for Podisus.

Three data sets (character matrices) were created for each genus: external charaters, male genitalic characters, and female genitalic characters. Characters were mostly bistate but some were multistate. Each of these matrices was run 10 times and the resulting trees compared. In these cases, the "mid" option was not specified, causing the trees to be rooted between the outgroup and the species under consideration. The trees derived from each data set were more consistent for Apateticus, whereas for Podisus 22

many homoplasious characters (those showing reversals or

Parallelisms) were evident, particularly in the female

genitalic data set. This shows the very close relationship among the species of the subgenus Apateticus and especially

that of the subgenus Apoecilus. The results for Podisus

demonstrate the uniformity of the external and internal

female genitalia, a fact noted by McDonald (1966) for the Asopinae in general.

For the final cladistic analysis, all data sets for each genus were combined after homoplasious characters were deleted. Subsequent to the initial run of these 2 matrices, characters showing a consistency index

(indicating homoplasious steps) of 0.50 or less were eliminated (scale of 1 to 10). The resulting matrices were run 10 times and the final tree derived (see Appendix).

Characters and their states for the external traits, male genitalia, female genitalia, and final combined matrices are listed in the Appendix.

In considering the final tree, one should recall that only the Nearctic species of Podisus are included. Also, behavioral data, such as courtship and mating rituals, ecological characteristics, and biogeographical data were not included because they are virtually unknown for many species. 23

DISCUSSION OF TAXONOMIC CHARACTERS

All characters considered in this study are from adult individuals. In describing these I am including those mentioned in the species descriptions, keys, and cladistic analysis. Measurements are expressed as ranaes which encompass the variation in specimens I examined.

BODY LENGTH. This was measured from the apex of the tylus to the tip of the hemelytral membrane with the specimen in dorsal view. Females generally exceed males in size. This measurement is fairly reliable in separating

Apateticus from Podisus although some overlap may exist between larger Podisus species and smaller individuals of

Apateticus. Within each genus, body length is not always diagnostic for any given species. Much variation within species exists and is probably due in part to the number of prey items consumed during nymphal development (see

Evans 1982b).

HUMERAL WIDTH. This was measured between the lateral apices of the humeral angles, also in dorsal view. In species where the humeral spines curve anteriorly, measurement was made between the spines' tips. Again,

females are usually larger than males and intraspecific variation can exceed that between species. 24

COLOR. In Apateticus, members of the subgenus

Apoecilus are of relatively uniform coloration and distinct from the subgenus Apateticus. In the latter, color is useful in species separation. Nearctic Podisus, excepting

P. acutissimus Stal and P. mucronatus Uhler, are not reliably distinguished solely by their coloration. The former 2 species have distinctive color patterns on the scutellum that are diagnostic.

HEAD. Width was measured between the lateral margins of the eyes with the specimen oriented dorsally. Length was also measured dorsally between the tip of the tylus and the base of the head. Females are generally larger. Both measurements vary between genera but not too much among species. The antennae are quite variable both in segmental length and coloration.

THORAX. Pronotal shape varies according to humeral configuration. Some differences were found in the ratio of pronotal length anterior to the humeri to total pronotal length, but these are not species recognition characters. The anterolateral margins of the pronotum are distinctive in a few species (A. marginiventris (St'Al) P. placidus, P. congrex (St5.1), P. mucronatus) but are concave - dentate in most. In some species (P. congrex,

P. acutissimus, P. sagitta, P. mucronatus, P. placidus, and A. marginiventris) the humeri are diagnostic due to their shape. The scutellum is fairly uniform. Width was 25 measured between the inner borders of the basalfoveae while length refers to the distancefrom base to apex, measured centrally. Females are usually larger than males. The hemelytral membrane may or may notpossess a dark longitudinal band. Podisus placidus, P. mucronatus, all members of the subgenus Apoecilus, and Apateticus marginiventris lack a band. Legs alone do not provide useful diagnostic characters for most species, although they are useful in distinguishing the subgenus Apateticus.

A black band is present apicallyor both apically and basally in the latter.

ABDOMEN. Ventral maculation is not diagnostic unless used with other characters. Within a given species of Podisus, small black spots may be present, indistinct, or absent. The abdomen is immaculate in P. congrex and

P. mucronatus. Taken with other characters, for instance in P. fretus, this trait may be useful. Differences in punctation are not obvious except in P. mucronatus which has larger ventral punctures than other species. In conjunction with other traits, the ventral spine isa useful means of species recognition. I have described this structure as not reaching the metacoxae, extending between the metacoxae, or reaching the anterior border of the metacoxae. The ventral spine varies from slender to stout. 26

MALE GENITALIA. Terminology used is that of Baker

(1931) and McDonald (1966). The Asopinae possess the following genitalic characters which together separate them from the rest of the Pentatomoidea (McDonald 1966):

(1) Presence of a pair of genital plates attached to the dorsal margin inside the genital cup; (2) Membranous conjunctival appendages; (3) Presence of a thecal shield at the apex of the theca; (4) Presence of a pair of median penial lobes encircling the tip of the vesica; (5) A simple ejaculatory reservoir with the seminal duct entering a posterior canal. The endophallic duct and seminal duct enter the reservoir adjacent to each other. McDonald (1966) notes that the genitalia of male Asopinae are very uniform. I was unable to detect distinct or consistent interspecific differences in the aedeagus, and therefore have not relied on it for species separations.

In Apateticus the thecal shield is much larger than the theca, whereas in Podisus these 2 structures are more nearly equal in size. Additional differences are described below.

Genital cup width measurements were made on dissected cups in dorsal view between the outer borders of the apices of the lateral lobes. This varies between species but is fairly constant within. Genital cups were grouped into 3 types for the cladistic analysis. Type A has prominent, lateral lobes, 2 mesas setal tufts on the ventral border 27 which is subtended by a deep concavity, and is typical of the subgenus Apoecilus (figs. 3b, 4b, 5b). Type B has narrower lateral lobes than Type A. The area between the ventral border and the ventral surface of the genital cup varies from flattened and slightly concave to deeply concave. Type B is characteristic of the subgenera

Apateticus and Podisus (figs. lb, 2b, 6b, 8b, 9b, 10b,

11b, 12b, 13b, 14b). Type C, characteristic of P.

(Tylospilus) acutissimus, has lateroventrally protruding lateral lobes, a narrow dorsal surface, and ventral margin with a U- or V-shaped notch. This cup is tilted forward so its contents are obscured in the caudal view (fig. 7b).

The proctiger is distinctive in some species of Apateticus, and Podisus due to a median projection (fig. 3b, 4b) or ridges (fig. 6c), but otherwise rather uniform.

Clasper shape is the most useful single diagnostic character at the species level. Alone it suffices to separate all but A. marainiventris from A. lineolatus and

P. maculiventris from P. serieventris. Again, for cladistic analysis, the claspers were arranged into 5 different types. Type A, seen in the subgenus Apoecilus, is bifurcate with specific differences in relative thickness, length, and shape of the dorsal and ventral arms (figs. 3c, 4c, 5c). Type B, found in the subgenus Apateticus, is thin, bladelike, tapered to a point, and curved dorsolaterad (figs. lc, 2c). Type C occurs in all 28

species of the subgenus Podisus except P. mucronatus and

is triangular with varying degrees of lateral indentation which creates differences in shape in the upper and lower

portions (figs. 8c, 9c, 10c, 11c, 12c, 13c, 14c).Type D

is found in P. mucronatus and the outgroup S. cincticeps.

This clasper is V-shaped with pointed apices (fig. 6c).

Type E, represented by P. acutissimus, is a flat, broadly and bluntly sickle-shaped clasper oriented so the flat sides face each other when the insect isviewed caudally

(fig. 7d).

The genital plates are not reliablespecies diagnostic characters. Apateticus spp. have plates shaped like an inverted L (figs. la, 2a, 3a, 4a,5a); the width of the stem andbase vary. Podisus spp.genital plates are quadratewith rounded corners andindented or bulging margins (fig. 15j). The single pair of conjunctival appendages differs between genera. Those of the subgenus Apoecilus have sclerotized processes dorsally

(fig. 15e), while in the subgenus Apateticus, only the tips are sclerotized (fig. 15f). In Podisus, these structures are entirely membranous. These structures are not useful in separating species.

The median penial lobes surrounding the tip of the vesica are long and tapered in the subgenus Apoecilus

(fig. 15g) and disk-shaped in the subgenus Apateticus and the genus Podisus (fig. 15h). These structures are also 29 of no real value in species distinction.

FEMALE GENITALIA. Terminology used is that of

McDonald (1966) and Schaefer (1968). Most structures are

very uniform and cannot alone distinguish species. The

external genitalia are virtually identical within the

subgenus Apateticus and the genus Podisus. The first

gonocoxae (first valvifers) are basically quadrate and

often margined mesally or posteriorly with brown. In the subgenus Apoecilus the triangulin (fused first valvulae),

a structure located between the first gonocoxae and attached to their anterolateral corners, is exposed and

differs in shape among the species. This structure is

covered by the first gonocoxae in the subgenus Apateticus and the genus Podisus.

McDonald (1966) studied 12 genera of Asopinae and noted the constancy of the spermatheca. This structure consists of a median dilation with a sclerotized rod

running lengthwise down the middle. A duct passes down the center of the rod and emerges from the dilated area to

a pumping region that is characterized by proximal and distal flanges. The spermathecal bulb is attached to the pumping region.

The right first gonocoxa was measured after clearing the abdomen in 10% KOH. Width refers to the distance between the lateral and mesal corners along the caudal border. The length was measured along the mesal margin 30

between the anterior and posteriorcorners. In the

subgenus Apoecilus, the shape of the firstgonocoxae and

triangulin are good species recognition characters. (The triangulin has also been called the median basal plate

(Heidemann 1904; Furth 1974; McPherson 1982).) The spermathecal duct varies in width and configuration between the 2 genera. In Apateticus, the duct is wide between the spermathecal dilation and the pumping region. It may be straight or coiled, dependingon the species (figs. le, 3d,

4d). Podisus has a narrow duct which is not coiled.

The spermathecal bulb was the most reliable female genitalic character. APateticus, excluding A. marqiniventris, has a characteristically small, elliptical bulb with prominent flanges (figs. le, 3d, 4d, 5d). P. placidus and P. saqitta can be distinguished by their round and fusiform bulbs, respectively (figs. 13e, 14e). For other species, differences exist but are not consistent enough to be reliable. Scierotized rod measurements vary considerably and are not useful diagnostic characters. 31

TAXONOMY

The introductions to Apateticus Dallas and Podisus

Herrich - Schaeffer are presented together rather than

preceding each genus because their history is very

intertwined. The genus Apateticus was erected by Dallas in 1851 to contain his newly described Apateticus halys.

In 1853 Herrich - Schaeffer created Podisus into which he placed no species until 42 pages later in the same work

(Kirkaldy 1908). Here Herrich - Schaeffer described

Podisus punctipennis, P. striqipes, P. vittipennis, P. pallipes, and P. albiseptus.

atIll in 1860, erected the genus Telepta into which he placed some species formerly contained in Arma Hahn as well as some newly described species. In 1867 St&l, apparently overlooking Apateticus as the older name, divided the genus Podisus into 4 subgenera: Telepta,

Troilus, Apateticus, and Podisus. In the first general work on Podisus, St&l (1870) rearranged the genus to include 5 subgenera: , Apateticus, Apoecilus,

Podisus, and Tylospilus, sinking Telepta as a synonym of the subgenus Podisus. He transferred Podisus punctipennis to the subgenus Apateticus, P. striqipes to Mineus St&l, P. albiseptus to Tvnacantha Dallas, and left P. pallies 32 as uncertain. Podisus (Troilus) luridus (F.), the only species in its subgenus and the sole Old World Podisus, was given generic status by StIl in 1872. Kirkaldy (1909) transferred this species to the genus Arms.

Distant (1880) followed StIl's generic classification, but did not designate subgenera, and described 11 new species from Central and South America between 1880 and

1893. Included in this work are color illustrations drawn from the type specimenwhere possible.

Schouteden (1907), noting that Apateticus preceded

Podisus, placed all species into the former. He divided the genus into the same subgenera as St&l, substituting the name Eupodisus for Podisus, and fixed type species for each. Kirkaldy (1909) followed Schouteden but used the subgenus Podisus rather than Eupodisus, and fixed P. vittipennis H.-S. (=Arma niarispina Dallas 1851) as the type species of Podisus. Kirkaldy (1909) listed 68 species under Apateticus:

57 in the subgenus Podisus, 7 in the subgenus Tylospilus, and 2 each in the subgenera Apateticus and Apoecilus. Four species have since been added to the subgenus Podisus

(Olsen 1916; Barber and Bruner 1932; Barber 1939; DaCosta

1957), 1 to Tylospilus (HorvAth 1911), and 1 to the subgenus Apoecilus (Van Duzee 1935).

In the United States several workers have studied

Apateticus and Podisus taxonomically. Say (1831), Uhler 33

(1870, 1871, 1897a, b), Fitch (1856), and Van Duzee (1935)

all described species in these genera. Kirkland (1898) published on the U. S. species of Podisus, including what are now considered Apateticus, and noted their value as predators.

Van Duzee (1904) provided an annotated list of Podisus

including Apateticus as a subgenus. In 1909, he took issue with Schouteden's classification and suggested that

Apateticus and Podisus be considered separate genera with the subgenera Apateticus and Apoecilus under the former and Podisus and TylosPilus in the latter. Van Duzee's

(1917) Catalogue of the Hemiptera is the most recent literature compilation and listing of North American

Podisus and Apateticus. Included are 14 species.

I have maintained Van Duzee's (1909) separation of the genera for the following reasons: (1) Differences in male and female external and internal genitalia, discussed in the section on taxonomic characters, p. 23; (2) The overwintering stage of Apateticus is the egg, while in

Podisus adults overwinter (p. 3). Unlike Van Duzee

(1909, 1917) I have not included P. fuscescens (Dallas) and treat P. concfrex (St&l) as an incidental Nearctic species. 34

GENUS APATETICUS DALLAS

Apateticus Dallas, 1851: 105. Type species = Apateticus

halt's Dallas 1851 (synonym of Halys lineolata H.- S.

1839); by monobasy. Schouteden (1907: 70) also fixed

this species as the type, as Apateticus lineolatus.

Podisus, subgenus Apateticus Stal, 1867: 498.

Podisus, subgenera Apateticus and Apoecilus Stal 1870: 48. Type species of subgenus Apoecilus = Pentatoma cynics

Say 1831 (synonym of Arma qrandis Dallas 1851); by

subsequent designation (Schouteden 1907: 70, as Apateticus cynicus).

Podisus, in part, Distant 1880: 36; Kirkland 1898: 413. Apateticus, subgenera Apateticus and Apoecilus Schouteden

1907: 70; Kirkaldy 1909: 22; Van Duzee 1917: 76.

DIAGNOSIS: Longer than 12mm; juga convergent anteriorly.

Males with rugulose, setose areas on sternites IV - VI;

fore femora unarmed; fore tibiae not dilated. Claspers bifurcate or falcate; thecal shield much larger than theca;

conjunctival appendages with sclerotized processes dorsally

or sclerotized at tips only. Females with triangulin exposed or covered by first gonocoxae.

DESCRIPTION: Body length: male 11.5 - 19.0; female 12.5 -

21.0. Humeral width: male 6.1 - 10.2; female 6.4 - 17.5.

Shape ovate-elongate, tapered behind middle; in females of 35

some species abdomen may widen at middle. Dorsum

ochraceous to brown, mottled grayish yellowor yellowish brown, at times with darkgreen cast; venter pale yellow

or shades of brown sometimes with brassy tint. Dorsal

punctation darker than ground color, most denseon juga,

pronotum, scutellum; venter more finely punctate than dorsum. Head length and width subequal; sides sinuate

anterior to eyes; juga exceeding tylus, margins convergent

apically; tylus with thickened margins, each subtended mesally by short sulcus. Rostrum stout, reaching hind coxae. Antennae about half body length, rufous to fuscous or ochraceous, finely setose, second segment usually

longest. Pronotum declivous cephalad of humeri; antero- lateral margins either concave-dentate or straight- edentate. Humeri blunt to acute or produced. Scutellum longer than wide, foveate in each basal angle; frena exceeding middle; apex concolorous or slightly paler than rest; often with irregular, pale, median longitudinal line.

Hemelytra tinged with red ventrally on embolium and apex or concolorous. Membrane clear to brown, without dark spots (except in A. lineolatus). Legs usually concolorous with venter, finely punctate, tibiae with 1 or 2 black bands or unmarked; fore tibiae not dilated; fore femora without spine or tubercle. Venter immaculate (except in

A. lineolatus); males with hairy rugulose area on each side of midline of sternites IV - VI. Ventral spine thick, 36

blunt, reaching anterior margin of hind coxae, or thin to very small. Duct of metathoracic scent gland extending laterally 1/2 to 2/3 width of metapleuron, gently curved

forward in apical 1/2. Genitalia: male genital cup with prominent, broadly rounded to tapered lateral lobes; claspers bifurcate or falcate. Female with triangulin exposed or covered by first gonocoxae; spermathecal duct between dilation and pumping region wide, looped or straight; bulb small, elliptical to oval; flanges prominent.

DISTRIBUTION: New World. Three species, A. bracteatus

(Fitch), A. anatarius Van Duzee, and A. cynicus (Say) occur only in the Nearctic region. A. lineolatus (H.-S.) and A. marqiniventris (St&l) have a Nearctic - Neotropical distribution.

Key to the Species of Apateticus

1. Humeri blunt, rounded (figs. la, 2a); ventral spine

short and thin or bumplike (fig. 2f); dorsum mottled

grayish yellow or yellowish brown with darker

overtones; claspers falcate (figs. lc, 2c); female

with triangulin covered by first gonocoxae (subgenus

Apateticus) 2

Humeri subacute to acute (fig. 3a, 4a); ventral spine stout, blunt, reaching anterior margin of hind coxae

(fig. 5a); dorsum ochraceous to sienna, sometimes 37 overlaid with dull black; claspers bifurcate (figs. 3c,

4c, 5c); females with triangulin exposed (figs. 3e,

4e, 5e) (subgenus Apoecilus) .. 3 2. Anterolateral margins of pronotum concave, dentate;

dorsum mottled grayish yellow with fuscous punctation; ventral spine thin, extending between metacoxae

* ***** ...... OOOOOOOOOO . OOOOO ...... lineolatus, p. 38

Anterolateral margins of pronotum, basal 1/2 of

hemelytral margin pale yellow; dorsum yellowish brown with gray or dark green cast; ventral spine reduced to

small swelling marqiniventris, p. 43

3. Clasper with dorsal arm bent caudolaterad at

approximately 90°in apical half; both arms blunt

apically and of equal thickness (fig. 4b, c); female triangulin with V-shaped ridge (fig. 4e); species

known only from Arizona anatarius, p. 56 - Clasper with dorsal and ventral arms parallel or

subparallel 4

4. Clasper with dorsal and ventral arms parallel, the

former 2/3 the length of the latter, slightly thinner

(fig. 3c); female triangulin triangular (fig. 3e)....

* **0***04 OOOOOOOOOOOOOOOOOOO Ot*O00** bracteatus p. 47

- Clasper with thin dorsal arm parallel or subparallel,

extending half the length of blade-like ventral arm

(fig. 5c); female triangulin quadrate (fig. 5e)

cynicus, p. 59 38

Subgenus Apateticus Dallas

DIAGNOSIS: Humeri bluntly rounded; tibiae with 1or 2 black bands. Genital cup with lateral lobes tapered at

apex (not broadly rounded); claspers falcate; median penial

lobes rounded, disk-like; conjunctival appendages with

sclerotized tips. Female triangulin covered by first

gonocoxae; shape of spermathecal duct and bulb characteristic.

Apateticus lineolatus (Herrich- Schaeffer)

(Figs. 1, 5c, f; Map 1)

Halys lineolata Herrich- Schaeffer 1839: 69, fig. 514. Apateticus halys Dallas 1851: 105, pl. 2 fig. 2.

Podisus (Apateticus) halys Stal 1870: 49.

Podisus punctipennis Herrich- Schaeffer 1853: 338.

Podisus (Apateticus) lineolatus Stal 1872: 129; Distant

1880: 36, pl. 1 fig. 39.

Apateticus (Apateticus) lineolatus Schouteden 1907: 70;

Kirkaldy 1909: 22; Van Duzee 1917: 76.

TYPES AND TYPE LOCALITIES: According to Dr. M. Baehr (pers.

comm.), Zoologische Staatssamlung, Munich, West Germany, Herrich - Schaeffer's types, Halys lineolata and Podisus punctipennis were probably destroyed in World War II. 39

DIAGNOSIS: Color mottled brown to gray; tibiae with black basal and apical bands; humeri blunt; anterolateral borders of pronotum dentate.

DESCRIPTION: Body length: male 12.5 - 14.5; female 14.0 -

17.5. Humeral width: male 6.1 - 7.2; female 6.7 - 8.7. Dorsum mottled gray to brown, basal half of pronotum, hemelytra pale grayish yellow with fuscous punctation; head, anterior half of pronotum, scutellum orange - yellow with reddish brown punctation; venter pale yellow to yellowish orange. Head with juga widely bordered in black; 2 black lines mesad oe each eye; margins slightly reflexed; width: male 2.5 - 2.8; female 2.7 - 3.1; length: male 2.4 - 2.8; female 2.6 - 3.2.Antennae fuscous with pale incisures; segment I pale yellow dorsally, black below; II paler fuscous than III, IV, V.

Pronotum with concave anterolateral borders denticulate on anterior half; scattered clusters of black punctures;

2 raised yellow to orange spots abutting posterior border of each callus. Humeri broadly rounded, not produced. Posterolateral margins of pronotum sinuate.

Scutellum: width: male 3.2 - 3.7; female 3.4 - 4.5; length: male 4.0 - 5.0; female 4.2 - 5.8; 3 - 5 small pale spots basally, separated by black areas; irregular pale longitudinal median line extending from vertex of head to apex of scutellum. Membrane with dark spots. 40

Legs yellowish to orange; femora mottled with fuscous; tibiae with apical and basal fuscous bands; tarsi fuscous. Venter of female with fuscous ring-shaped spot on sternite

VI. Male genitalia: width of genital cup 1.91 - 2.17; proctiger subvertical with apical ridge subtended by flattened black area; cup semicircular in caudal view; lateral lobes ending in bluntly rounded point; area between ventral border and ventral surface of cup deeply excavated, black with ventral setal tufts; claspers slender, falcate, directed dorsolaterad; genital plates with longitudinal lines or grooves; dorsal margin with deflexed rounded tubercles. Female genitalia: width of first gonocoxa 0.80 - 0.95, length 0.66 - 0.80; spermathecal duct between dilation and proximal flange long, bent, coiled; length of scierotized rod 0.75 - 1.21.

SPECIMENS ILLUSTRATED: Humerus, head, fuscous ring-shaped spot of female, gonocoxae: Texas: Uvalde Co., Concan

(AMNH); genital cup: Texas: Cameron Co., Esprza Ranch,

Brownsville (CU); clasper, conjunctival appendages

Texas: Cameron Co., Brownsville (JE); spermatheca:

Texas: Cameron Co., Brownsville (OHS).

DISTRIBUTION: Southern Texas, throughout Mexico, Central

America, and northern South America (Map 1). 41

MATERIAL EXAMINED: 200 specimens from the following

localities: COSTA RICA: Cartago: Turrialba (AMNH);

Pacayas (AMNH). Puntarenas: Monteverde (TAM). GUATEMALA:

Baja Verapaz: Rabinal (AMNH); San GerOnimo (AMNH).

MEXICO: Aguascalientes: Aguascalientes (UNL); Rinccin de

Romos 6100' (UMI). Chiapas: 21 km W Rizo de Oro (CAS); 4 mi W San Cristobal de las Casas (UMI); 17 km SE CristObal

de las Casas (CAS); "Chia." (UAZ). Coahuila: La Gloria,

S of Monclova (AMNH). Guanajuato: 64 km SE La Sauceda (DBT). Guerrero: 10 mi SW Coatlan del Rio (UMI); Taxco

(OSU); Telolcapan (UMI). Hidalgo,: 12.7 rd mi N Atotonilco

El Grande (N of Pachuca) (UMI); 15 mi S Jacala (JE); 15 mi

NE Huichapan (UMO). Jalisco: 16 km N Autlan (TAM);

Guadalajara (TAM); Puerto los Mazos, 9 mi NW Autlan (UCB);

Sierra Autlan, 20 mi SSE Autlan (UMI). Mexico: Bejucos,

Temescaltepec (UCB); Real de Arriba, Temescaltepec (UCB);

Tejupilco, Temescaltepec (UCB). Michoacan: Cerro San

Andres, 11 mi WNW Ciudad Hidalgo (UMI). Morelos:

Cuernavaca (CAS, ASU); 5.1 mi E Cuernavaca (TAM); 10 mi E

Cuernavaca (UCB); 3 mi NW Cuernavaca (CU). Nayarit:

Arroyo Santiago nr Jesus Maria (UCB); Cerro de San Juan,

4.5 rd mi SW Jalisco (UMI); Tepic (AMNH). Nuevo Le6n: 7.5 mi S Monterrey (JE). Oaxaca: 2.7 mi NW El Carmen (TAM);

Oaxaca 5034' (AMNH); 56 mi NW Tehuantepec (UCB); Oaxaca

(OSU); 16.1 mi NW Totolapan (JE). Puebla: 7.7 mi SE

Izticar de Matamoros (UMI); Matamoros (UMI); 17 mi S Puebla 42

(OSU). Queretaro: 1 mi NW Ayutla (TAM). San Luis

Potosi: 6 mi E Ciudad del Maiz (TTU); 52 km E Ciudad del

Maiz (UAZ). Sinaloa: Mazatlan (CAS); 2 mi E Tecorito, E

of Humaya (13.1 mi N Culiacan) (UMI). Sonora: Cyn Sapopa

(CAS); Sonora (AMNH). Tamaulipas: Abasolo (AMNH);

Bocatoma, 7 km SSE GOmez Farias (DBT); 6 mi S GcSmez Farias

(DBT); Grutas de Quintero, 8 mi S Cd Mante (UGA); 13 km E

Magiscatzin (UAZ); Rio Corona, 30 km NE Ciudad Victoria

(DBT); Victoria (AMNH). Veracruz: Atoyac (AMNH); 5 km W

Conejos (JE); C6rdoba (AMNH); Orizaba (CAS).Not located:

Mexico (AMNH, ISU). PANAMA: El VolcAn de Chiriqui (AMNH);

Portrerillos (ISU). UNITED STATES: Texas: Brewster Co.:

Chisos Mts (UID, OHS); Cameron Co.: Brownsville (AMNH,

CAS, OHS, JE, UCT); Brownsville - Esprza Ranch (CU); Sabal

Palm Grove Sanctuary (DBT, UGA); Cameron Co. (AMNH);

Dimmit Co. (TAM); Gillespie Co. (OHS); Goliad Co.: US 183,

6.7 mi S Weser (UAL); Hidalgo Co.: Bentsen - Rio Grande

Valley State Park (JE, DBT); Hidalgo Co. (OHS); Jeff Davis

Co.: Davis Mts (OHS); Real Co.: Camp Wood (TTU); Sutton

Co. (AMNH); Uvalde Co.: Concan (AMNH); Uvalde (CAS); Val

Verde Co.: Devil's River (OHS); 6 mi S Juno (TTU).

This species may be confused with the genus Brochymena

Amvot & Serville (Pentatominae), as both are mottled gray or brown. A. lineolatus has a stouter beak than Brochymena and the first rostral segment is not embedded between the bucculae. Species of the latter are generally flatter than 43 A. lineolatus.

A. lineolatus is found from southern Texas,an area equivalent to Bailey's (1978) Mesquite AcaciaSavanna section of the Prairie Brushland Province,throughout

Mexico, Central America, and into Venezuela. Habitat associations are unknown.

Plummer and Landis (1932) list this speciesas the second most abundant pentatomid predator of the Mexican bean beetle (Epilachna varivestis Mulsant) inCuernavaca, Morelos, Mexico. Collections were made in August and

September. They obtained one oviposition record of 93 eggs in a single mass. Their rearing results for 18 field collected nymphs observed in thelaboratory are presented in Table 1.

Apateticus marqiniventris (Sthl)

(Fig. 2; Map 2)

Podisus (Apateticus) marainiventris St&l 1870: 49.

Podisus marqiniventris Distant 1880: 36,pl. 4, fig. 24. Podisus qillettei Uhler 1895: 12; Kirkland 1898: 418; Van Duzee 1904: 69.

Apateticus (Apateticus) marqiniventris Schouteden 1907:70; Kirkaldy 1909: 22; Van Duzee 1917: 77.

TYPES AND TYPE LOCALITIES: Podisus (Apateticus) marqini- ventris Sth, type, female,very good condition (Riks- museet): "Oaxaca / Sa114. / Typus (red) / 412;82 (red) / 44

Riksmuseum; Stockholm (green)." Type locality is Oaxaca,

Mexico. Podisus qillettei Uhler has not yet beenlocated.

DIAGNOSIS: Anterolateral margins of pronotum, basal 1/4

of corium margined with pale yellow; humeri blunt;ventral spine reduced to small swelling.

DESCRIPTION: Body length: male 11.5 - 13.5; female 12.5 -

14.5. Humeral width: male 6.1 - 7.2; female 6.4 - 8.0.

Dorsum yellowish brown with a gray or darkgreen cast,

black punctation; venter yellow withorange tint, finely

punctured with red to fuscous. Head declivous, densely

punctate with black; width: male 2.5 - 2.8; female 2.6 -

3.0; length: male 2.2 - 2.5; female 2.4- 2.8. Antennae fuscous to piceous, incisures pale; segment I pale yellow,

fuscous ventrally. Pronotum declivous anteriorly; margins yellow, straight, edentate to slightly crenulatenear apices. Humeri blunt, broadly rounded. Scutellum: width: male 3.4 - 4.0; female 3.6- 4.5; length: male 3.9 - 4.5; female 4.1 - 5.2; densely punctate with blackon basal 2/3.

Membrane light brown.Legs variable brown to fuscous; coxae pale, tibiae black apically. Ventral spine very short, not reaching hind coxae; often a small, pointed swelling. Male genitalia: width of genital cup 2.01-

2.17; lateral lobes smaller,more rounded apically than preceding species; proctiger broadly rounded dorsally, setose tip reflexed; area between ventral border and 45 ventral surface flattened, as though latter were bent

abruptly dorsad; claspers slender, falcate, wider basally

than preceding species; genital plates very similar to preceding species. Female aenitalia: width of first gonocoxa 0.52 - 0.93; length 0.52 - 0.80; spermathecal duct thin, straight or looped between dilation and pumping

reaion; bulb rounded to pyriform, larger than others in genus; length of sclerotized rod 0.88 - 1.72.

SPECIMENS ILLUSTRATED: Humerus, genital cup: Arizona:

Santa Cruz Co.: Patagonia (CAS); ventral spine: Arizona:

Santa Cruz Co.: W slope Patagonia Mts on Lochiel Rd 5330'

(UAZ); clasper: MEXICO: Sonora: El Tigre (UMI); spermatheca: Arizona: Santa Cruz Co.: Canelo (UAZ).

DISTRIBUTION: Montana, Colorado, Arizona, New Mexico; Chihuahua, Michoacgn, Oaxaca, and Sonora, Mexico (see Map 2).

MATERIAL EXAMINED: 24 specimens from the following localities: MEXICO: Chihuahua: 8 mi W Matachic (AMNH);

Matachic (AMNH). Michoacan: Pedregal, nr Tancltaro, 6000'

(AMNH). Sonora: Sierra de Madera, 5 mi E El Tigre (UMI).

UNITED STATES: Arizona: Cochise Co.: Huachuca Mts, 3 mi

E Montezuma Pass, 5000 - 5500' (AMNH); Huachuca Mts, Ramsey

Canyon (CAS, UAZ); Santa Cruz Co.: Canelo (UAZ); Palmerlee

(AMNH); Patagonia (CAS); W slope Patagonia Mts on Lochiel

Rd, 5330' (UAZ). Colorado: Larimer Co.: Fort Collins 46

(CAS, CSU). Montana: Gallatin Co. (ISU, MSU). New

Mexico: Santa Fe Co.: Tesuque (AMNH).

This species varies in the amount ofgreen or gray

overtones in its coloration. I observed very little

variation in the male genitalia andonly minor differences

in the spermathecal bulb. A. marqiniventris apparently occurs as very localized populations over a large

geographical area (Map 2). Its biology is unstudied and the nymphs and eggs have not been described.

Subgenus Apoecilus St&l

Podisus, subgenus Apoecilus Stal 1870: 48. Type species =

Pentatoma cynica Say 1831. (synonym of Arma qrandis

Dallas 1851); by subsequent designation (Schouteden 1907: 70, as Apateticus amigap).

Apateticus, subgenus Apoecilus Schouteden 1907: 70;

Kirkaldy 1909: 22; Van Duzee 1917: 70..

DIAGNOSIS: Coloration ochraceous; humeral angles subacute to acute; ventral spine stout, extending between hind

coxae. Genital cup with prominent, rounded lateral lobes; shelf-like ventral border with 2 tufts of setae curving

and meeting medially; claspers bifurcate; median penial

lobes of aedeagus long, gradually tapering to blunt apices; conjunctival appendages with sclerotizedprocess dorsal to membranous area. Female triangulin exposed; 47 spermathecal duct wide distad of dilation; bulb small, elliptical; flanges prominent.

Apateticus bracteatus (Fitch)

(Figs. 3, 15e, g; Map 3)

Arma bracteata Fitch 1856: 336.

Podisus crocatus Uhler 1897a: 384; Kirkland 1898: 422.

NEW SYNONYMY. Podisus (Apoecilus) bracteatus Van Duzee 1904: 70.

Apateticus (Apoecilus) bracteatus Schouteden 1907: 70; Van Duzee 1917: 78.

Apateticus (Apoecilus) bracteatusvar. crocata Kirkaldy 1909: 22.

TYPES AND TYPE LOCALITIES: Arma bracteata Fitch, type,

female, very good condition (USNM): "15,353 / Fitch's

Type / Fitch's Collection / Podisus (Arma) bracteatus

Fitch / Type No. 1019 U. S. N. M. (red)." No locality data

are given, but presumably taken in New York. Podisus crocatus Uhler, no type label, female, good condition

(USNM): "Colville V., W. T.; Brown's; July 24, '82 / P.R. Uhler Collection / Podisus crocatus Uhler; W. T.; Det.

Uhler / Podisus crocatus; W. T.: Uhler." Colville is in present - day Stevens Co., Washington. As this specimen bears no red label or type designation, I willpropose that it be given lectotype status in a subsequent paper. 48

DIAGNOSIS: Coloration orange - yellow to reddish orange; humeri subacute to acute. Clasper with arms parallel; dorsal arm shorter. Female triangulin triangular.

DESCRIPTION: Body length:male 12.5 - 16.5; female 13.0 -

18.0. Humeral width: male 6.4 - 8.8; female 6.9 - 10.0. Dorsum orange-yellow to reddish orange; punctation nearly concolorous to reddish brown or dull greenish black; venter pale yellow to oranae with fine red or brown punctation.

Head with juga of dark specimens often margined with areenish black; width: male 2.6 - 3.1; female 2.8 - 3.4; length: male 2.3 - 2.9; female 2.5 - 3.3. Antennae ochraceous to rufous or fuscous. Pronotum with antero lateral margins concave-dentate; usually darker on basal half which may be faint to dark greenish black; anterior half often with greenish black spots. Humeri subacute to acute; apices often black. Posterolateral pronotal margins sinuate, ending bluntly or in tiny acute tooth. Scutellum: width: male 3.2 - 5.4; female 3.7 - 5.0; length: male:

4.0 - 5.7; female 4.3 6.8. Legs concolorous with venter or rufous; femora densely punctate. Male genitalia: width of genital cup 2.25 - 2.97; proctiger with dorsomedian, deflexed projection; clasper with parallel arms, dorsal arm 2/3 length of ventral arm which ends in deflexed, acute point; clasper height 0.42 - 0.64; genital plates shaped like inverted L; margins finely tuberculate; tubercles scattered on surface of plate. Female genitalia: width of 49

first gonocoxa 0.80- 1.19; length 0.66 - 1.72; triangulin

triangular, often with small depression; spermathecalduct coiled or looped; length of sclerotized rod 1.22- 2.60.

SPECIMENS ILLUSTRATED: Humerus, genital cup: New York: Tompkins Co.: Ithaca (CU); clasper, conjunctival appendages: Utah: Davis Co.: Kina's Station (CAS); aedeagus, median penial lobes: Washinaton: Whitman Co.: Pullman (WSU); humerus, gonocoxae: CANADA: British

Columbia: Victoria (UBC); spermatheca: Montana:

Gallatin Co.: Gallatin Valley (MSU).

DISTRIBUTION: Transcontinental in the north; south to

California, New Mexico, Illinois, and Maryland (Map 3).

MATERIAL EXAMINED: 456 specimens from the following localities: CANADA: Alberta: Cypress Hills (UAL);

Edmonton (CAS, UMI, OSU); Medicine Hat (UAL); Milk River

(UAL); Rockyford (UAL); Wellington (UAL). British Columbia:

Eburne (UBC); Royal Oak (UBC); Saanich District (CAS, UBC);

UBC ForeSt (UBC); Vancouver (UBC); Vancouver Island (AMMH);

Victoria (INH, PDU, UBC); not located (UID). Manitoba:

Winnipeg (UAZ); not located (UAZ). Nova Scotia: Cape

Breton Island (CMNH); Truro (CU); Yarmouth (CAS). Ontario:

Cardinal (UMI); Inverhuron Park (INH); 15 mi SE Kenora

(UCB); Ridgeway (ISU); Swansea (ISU); Toronto (OSU).

Quebec: Covey Hill (UBC); Lauzon (TAM); Mon-real (CAS,

UBC). Saskatchewan: Lloydminster (UMO). UNITED STATES: 50

California: Eldorado Co.: Fallen Leaf Lake (CAS); Lake

Tahoe (CU); Fresno Co.: Huntington Lake (CAS); Lassen Co.:

9 mi WMacArthur (AMNH); 3 mi W Nubieber (AMNH); Modoc

Co.: Buck Creek (CAS); Mono Co.: Mammoth Lake (CAS);

Sonora Jct, 6850' (OSU); Plumas Co.: Meadow Valley (UCB);

Plumas Co. (DBT); San Bernardino Co.: Mill Creek Canyon

(CAS); San Diego Co.: Cuyamacha Lake, 5000' (AMNH); San

Mateo Co. (CU); Santa Barbara Co.: Santa Cruz Island

(LACM, YLU); Sunset Valley (UID); Santa Clara Co.: Palo

Alto (LACM); Santa Cruz Co.: Mt. Hermon (CAS); Shasta Co.:

Sacramento District, Shasta National Forest (UCB);

Siskiyou Co. (CAS); Sonoma Co.: S Sonoma Co. (CAS);

Trinity (UCB); Trinity Co.: Carrville (CAS); Coffee Creek

(UCB); 8 mi S Zenia (UID); Tulare Co.: Giant Forest (CAS);

Tuolumne Co.: Pinecrest Col. (CAS); not located: Baldy (LACM); Clarksburg (UCB); Descanso (CAS); Laguna Mt (CAS); 15 mi W Mineral (OHS); Paraiso Sprgs (CAS); Riverton (UCB);

Twin Rocks (CAS). Colorado: El Paso Co.: Colorado

Springs (AMNH, OHS, UCT); Manitou Springs (AMNH); Huerfano

Co.: East Spanish Peak (UMI); Larimer Co.: Fort Collins

(ASU, CAS, CSU); Las Animas Co.: Trinidad (UAZ); Montezuma Co.: Mesa Verde (UCB); Montezuma Co. (OHS); Montrose Co.:

Black Canyon National Monument (UAZ); Routt Co.: Steamboat

Springs (OSU); Teller Co.: Pike's Peak (CU); not located:

"Dtch Geo" (CSU); Little Beaver (UAZ). Connecticut:

Fairfield Co.: Brookfield (UBC); Litchfield Co.: 51

Litchfield (AMNH); New Haven Co.: S Meriden (CAS).

Idaho: Bear Lake Co.: nr Fish Haven (UCB); Boise Co.:

20 mi N Boise (OSU); Butte/Blaine Co.: Craters of the Moon

Natl Monument (UID); Cassia Co.: Elba-Basin Pass (UID);

Latah Co.: Moscow (CAS, ISU); Moscow Mt (ISU, LSU, UID,

WSU); Potlatch (ISU); Nez Perce Co.: 5 mi SE Lake Waha

(UID); Shoshone Co.: Wallace (UCB). Illinois: Cook Co.:

Chicago (UNL); Lake Co.: Wauconda (INH); McHenry Co.:

Wonder Lake, Marengo (INH); not located: Dubois (INH);

"N. Ill." (INH, PDU); Volo, Tamarack Bog (INH). Maine:

York Co.: Wells Beach (AMNH); not located: Hempswell

(AMNH); Maine (CAS); Southport (ANKH). Maryland:

Baltimore Co.: Baltimore (AMNH). Massachusetts:

Barnstable Co.: Woods Hole (AMNH); Berkshire Co.: Adams,

Greylock Mt, 3500' (AMNH); Essex Co.: Ipswich (CAS, CMNH);

Salem and vicinity (AMNH); Hampshire Co.: Amherst (CAS);

Middlesex Co.: Maiden (CSU); Sherborn (AMNH, CAS);

Nantucket Co.: Nantucket Island (AMNH, INH); Norfolk Co.:

Dover (AMNH); not located: Beach Bluff (CAS, UBC);

Faneuil (AMNH); Stony Brook Res (AMNH); Sunderland (CAS);

Truro (AMNH). Michigan: Alger Co. (MIS); Allegan Co.:

Plainwell (MIS); Berrien Co.: E. K. Warren Preserve, Sawyer Dunes (UMI); Herbert Dunes (UMI); Charlevoix Co.:

Boyne Falls (UMI); Susan Lake (UMI); Thumb Lake (UMI);

Cheboygan Co.: Douglas Lake (UMI); Cheboygan Co. (UMI);

Chippewa Co.: Pickford (WSU); Trout Lake (UMI); Emmet Co.: 52

Waugoshance Island (UMI); Houghton Co.: Calumet (AMNH);

Huron Co.: Sand Point (UMI); Tosco Co.: Oscoda (UMI);

Iosco Co. (UMI); Livingston Co.: E. S. George Reserve

(UMI); Luce Co. (UMI); Mackinac Co.: Cut River (UMI);

Naubinway (UMI); St.Ignace (UMI); Marquette Co.: Huron

Mts (UMI); Midland Co. (LSU); Oakland Co.:Milford (UMI);

Oakland Co. (UMI); Ogemaw Co. (UMI); Washtenaw Co.: Ann

Arbor (UMI); Cavanaugh Lake (UMI); First Sister Lake (UMI);

Hamburg (UMI); Manchester (UMI); Washtenaw Co. (UMI);

Wayne Co. (UMI). Minnesota: Clay Co.: 2 mi N Comstock (UCB); Clearwater Co.: Upper Rice Lake (WSU); Freeborn

Co.: Albertlea (LSU); St. Louis Co.: Floodwood (CAS); not located: "Minn." (CU). Montana: Flathead/Glacier

Co.: Glacier Natl Park (MSU); Gallatin Co.:Gallatin

Valley (MSU); Gallatin Co. (MSU); not located: "Western

Montana" (MSU). Nebraska: Red Willow Co.: Indianola

(UNL). Nevada: Douglas Co.: Carson City (CAS); Washoe

Co.: Spooner Pass (OSU). New Hampshire: Grafton Co.:

Franconia (AMNH); not located: Crawfords (CAS). New

Jersey: Bergen Co.: Wyckoff (AMNH). New Mexico:

Sandoval Co.: Jemez Springs (NDSU); Santa Fe Co.: Santa

Fe (AHNH). New York: Erie Co.: Buffalo (CU, INH); Colden (ISU); Hamburg (CAS, ISU); Lancaster (AMNH, ISU,

UCR); Essex Co.: Moriah Center (OSU); Greene Co.: Rt

10, Town of Durham (UM0); Kings Co.: Flatbush (AMNH);

Livingston Co.: Dansville (UM0); Onondaga Co.: Syracuse .53

(LSU); Queens Co.: Rockaway Beach (AMNH); Suffolk Co.:

Fire Island (CU, PDU); Smith Pt LSS, Fire Island (CU);

Tompkins Co.: Ithaca (CU); not located: Maspeth (AMNH);

NY (AMNH, CU, ISU). North Dakota: Bottineau Co. (UMI);

Cass Co.: Fargo (NDSU); Cavalier Co. (NDSU); Divide Co.

(NDSU); Hettinger Co. (NDSU); Stutsman Co. (NDSU). Ohio:

Lake Co.: Willoughby (AMNH); Lake Co. (UCT); Lucas Co.:

Dorr & Cressy Rds (UMI); 6 mi W Toledo (UMI); Williams Co.

(YLU). Oregon: Benton Co.: Corvallis (ISU, LSU, OSU);

MacDonald Forest (OSU); Clackamas Co.: Oak Grove (OSU);

Clatsop Co.: Gearhart (OSU); Seaside (ISU, OSU, UCB);

Deschutes Co.: North Sister snowfields, 9000' (OSU);

Harney Co. (CAS); Jackson Co.: Colestin (CAS); Medford

(OSU); Siskiyou (CAS); Siskiyou Summit (CAS); Josephine

Co.: 10 mi E Cave Jct (GMS); Klamath Co.: Klamath Fails

(CAS); Lake Co.: Hart Mt, 6250' (AMNH); Lane Co.: Black

Butte (OSU); Marion Co.: Silverton Hills (OSU); Union Co.:

La Grande (OSU); Wasco Co.: Mt Hood (CAS); not located:

Springflats Camp, Scott River (WSU). Pennsylvania:

Crawford Co.: Hartstown (CMNH); Erie Co.: Lake Leboeuf

(CMNH). South Dakota: Brookings Co.: Brookings (CAS); not located: Lake Hendricks (ISU); "S. D." (CAS). Utah:

Cache Co.: Logan Canyon (AMNH); Davis Co.: King's

Station (CAS); Juab Co.: 4 mi E Nephi on Hwy 11, 6200'

(GMS); Weber Co.: Ogden (CAS). Washington: Asotin Co.: 54

Clarkston (OHS); King Co.: Donald, 1080' (ISU); Seattle

(OSU, WSU); Vashon Island (CAS);Mason Co.: Shelton (OSU);

Pacific Co.: Bay Center (OSU); Pierce Co.: Mt Rainier (ISU, LSU, WSU); Stevens Co.: Loon Lake (UMI); Wawawai (UCT); Whitman Co.: Pullman (ISU, KU, UCT, WSU); Yakima

Co.: Yakima (WSU). Wisconsin: Bayfield Co.: Bayfield

(ISU); Dane Co.: Madison (UWI); Door Co.: Bailey Harbor

(UWI); Iron Co.: Mercer (UWI); Marathon Co.: Wausau (YLU); Milwaukee Co.: Milwaukee (CU).

Kirkland (1898) placed this species (as Podisus crocatus) as a synonym of A. cynicus (Say) which it

resembles externally. He also mentioned Uhler's and

Glover's opinion that A. bracteatuswas a variety of A. cvnicus. No mention is made of the genitalia of either

species. Heidemann (1904) called attention to differences

in the male and female genitalia. Van Duzee (1904)

disagreed with Kirkland andgave A. bracteatus specific rank (as Podisus).

Van Duzee (1904) felt that A. crocatus wasa paler,

southern form of A. bracteatus but did notcare to

synonymize them until more materialwas available for study, When Uhler described A. crocatus he apparently did not know about bracteatus, for he allied his species with

A. cynicus. Downes (1920) believed that crocatus and bracteatus could be separated on the basis of bracteatus' more spinose humeri. This character is fairly consistent, 55

but both types of humeri can be found in specimensfrom Montana, Colorado, Utah, Illinois, North Dakota,

Massachusetts, New Hampshire, and Nova Scotia.Male and female genitalia are both inseparable and therefore I have synonymized the 2 forms.

A. bracteatus varies in color and the degree of

acuteness of the humeri over its geographicalrange. I detected very little variation in male and female

genitalia. The clasper shows some differences in the

thickness of the dorsal and ventralarms and amount of curvature of the former. The female genitalic structures are very constant except for minor differences in the

spermathecal bulb and coiling of the spermathecal duct.

Downes (1920) described the eggs and nymphs of this

species and reared it in the laboratory. Results are

given in Table 1. A. bracteatus is a polyphagous predator;

prey records are listed in Table 2. The species overwinters in the egg stage.

Ruckes (1938) collected A. bracteatus on Gambel's oak

(Quercus qambelii Nuttall) incanyons and at higher elevations around Santa Fe, New Mexico.This species was commonly taken along with Dendrocoris contaminatus Uhler

(Pentatomidae: Pentatominae) (Ruckes 1938). Stoner (1925) collected A. bracteatus on willows in July in the Douglas

Lake region in Michigan. I have taken it on willow in Oregon. 56

Evans and Root (1980) found A. bracteatus nymphs in

Solidaao sp. stands to be hunters which cluster in groups of 16 - 50 before molting, a process which occurs over 24

48 h. If disturbed, clusters would re-group within a short time. Older nymphs congregated in smaller groups.

Relationships among nymphs in a given cluster were not assessed, but Evans and Root (1980) suggested that most group members were siblings, owing to distance between aggregations. The authors also postulated a defensive role, as odors emitted en masse from dorsal abdominal glands and aposematic coloration (nymphs are red and black) could protect vulnerable nymphs from predation.

Percy et al. (1980) studied scent gland ultra- structure in A. bracteatus nymphs and identified tridecane and 2-octanal as major scent components. These chemicals are found in other nymphal pentatomid scents as well as adult metathoracic gland secretions (Staddon 1979).

Apateticus anatarius Van Duzee

(Fig. 4; Map 4)

Apateticus anatarius Van Duzee 1935: 27.

TYPE AND TYPE LOCALITY: Apateticus anatarius Van Duzee, holotype, male, excellent condition (CAS): "San Cata Mt.;

7-23-32; E. D. Ball. Ar. / HOLOTYPE; anatarius (red)."

Allotype, female, very good condition (CAS): "San Cata 57

Mt; 7-23-32; E. D. Ball. Ar. / 88 (on reverse) / ALLOTYPE; anatarius (red)." The type locality is the Santa Catalina

Mountains, Pima Co., Arizona.

DIAGNOSIS: Clasper with dorsal arm bent abruptly caudolaterad. Triangulin with V-shaped ridge.

DESCRIPTION: Body length: male 14.6 - 16.4; female 16.4 -

19.0. Humeral width: male 8.1 - 9.1; female 8.7 - 12.0.

Dorsum ochraceous to light brown with reddish brown punctation. Head with densely punctate juga; area between inner margin of each eye and imaginary extension of medial margin of juga with 3 longitudinal rows of brown to metallic green punctures; width: male 2.9 - 3.0; female

3.2 - 3.5; length: male 2.6 - 3.0; female 2.8 - 3.4.

Antennae ochraceous to rufous; apical 2/3 of segments III,

IV, and all of V darker. Pronotum with slightly concave anterolateral borders; anterior half dentate; disk somewhat rugose, some punctures metallic green. Humeri subacute, apically metallic green to black. Posterolateral pronotal margins slightly sinuate, each ending in small acute tooth which projects onto corium. Scutellum width: male 4.0 -

4.5; female 4.3 - 5.3; length: male 5.1 - 5.9; female

5.4 - 6.7. Legs concolorous with venter, covered with fine setae of mixed lengths. Male genitalia: width of aenital cup 2.92 - 3.02; proctiger vertical with beak-like projection set in obovate depression; dorsal arm of 58 clasper bent caudolaterad at approximately 900, extending half length of ventral arm; blunt apically, clasper height

0.64 - 0.80; genital plates shaped like inverted L, tuberculate marginally, sparsely so on surface. Female genitalia: width of first gonocoxa 0.95 - 1.06; length

0.85 - 0.95; triangulin with V-shaped ridge; spermathecal duct wide, straight between dilation and proximal flange; length of sclerotized rod 1.47 - 2.01.

SPECIMENS ILLUSTRATED: Humerus, ventral spine: Arizona:

Pima Co.: Catalina Mts (AMNH); genital capsule: Arizona:

Pima Co.: Catalina Mts (AMNH); clasper: Arizona: Santa

Cruz Co.: Santa Rita Mts (OHS); spermatheca, external genitalia: Arizona: Santa Cruz Co.: Santa Rita Mts (OHS).

DISTRIBUTION: Known only from several mountain ranges in southeastern Arizona (Map 4).

MATERIAL EXAMINED: 21 specimens from the following localities: UNITED STATES: Arizona: Cochise Co.:

Chiricahua its (CAS, DET, OHS); Huachuca Mts, Upper Carr

Canyon (CAS); Graham Co.: Pinaleno Mts (OHS); Pima Co.:

Catalina Mts (AMNH); Catalina Mts., Bear Wallow (AMNH);

Santa Cruz Co.: Santa Rita Mts (CAS, OHS); Santa Rita Mts,

Madera Canyon (CAS).

This species is known only from southeastern Arizona which, according to Bailey's (1978) ecoregions designations, 59 lies in the Mexican Highlands Shrub Steppe Province.

Bailey (1978) lists 4 life zones in this province: desert belt, arid grassland, submontane, and montane. Locality data for A. anatarius indicate the species' occurrence in mountains and canyons, most likely in the latter 2 of

Bailey's (1978) life belts. The species could be limited by unfavorable surrounding habitats or, conversely, it could be a relatively newly evolved species which has not yet spread past its area of origin. Also plausible is the notion that A. anatarius is a dying species. Before any of these speculations can be borne out, extensive collecting must be done.

The biology of this species has not been studied.

Apateticus cynicus (Say)

(Fig. 5; Map 5)

Pentatoma cynica Say 1831: 3.

Arena qrandis Dallas 1851: 96.

Podisus (Apoecilus) grandis Stal 1870: 49.

Podisus cvnicus Uhler 1876: 282; Kirkland 1898: 425.

Apateticus (Apoecilus) cynicus Schouteden 1907: 71;

Kirkaldy 1909: 22; Van Duzee 1917: 77.

TYPES AND TYPE LOCALITIES: The type(s) of Pentatoma cynica

Say, collected in "MiSsouri," is destroyed. I will propose a neotype from the type locality in a subsequent 60 publication. The type of Arma grandis-Dallas, in the

British Museum has not yet been seen.

DIAGNOSIS: Clasper with thin, short dorsal arm; ventral arm wider, bladelike. Female triangulin quadrate.

DESCRIPTION: Body length: male 14.0 - 19.0; female 16.0 -

21.0. Humeral width: male 7.6 - 10.2; female 7.8 - 12.0.

Dorsumvarious shades of brown, at times with bronze or orange overtones; punctation even, fine, brown; venter light yellow to light brown with brown to rufous punctation.

Head as in previous species; width: male 2.8 - 3.4; female 2.9 - 3.5; length: male 2.6 - 3.4; female-2.8'-

3.5. Antennae concolorous with dorsum or darker, sometimes rufous; apical half of segment IV, all of V usually darker than I - III. Pronotum with concave anterolateral borders; irregularly dentate anterior 1/2 to 2/3; 2 unequally sized groups of metallic green to black spots lateral to each callus. Humeri produced laterally, acute. Posterolateral pronotal margins straight to sinuate, ending in bluntly rounded corners. Scutellum: width: male 3.3 - 4.8; female 3.9 - 5.3; length: male 4.8 - 6.8; female 5.0 -

7.2. Legs colored as venter; femora with fine punctation; setose. Male genitalia: width of genital cup 2.22 - 2.65; proctiger without distinct beak-like projection; clasper with thin, short dorsal arm, sub - parallel to much wider blade-like ventral arm; clasper height 0.37 - 0.40; 61 genital plates shaped like inverted L, lateral margins rounded, edges and interior tuberculate. Female genitalia: width of first aonocoxa 0.74- 0.93; length 0.80 - 0.90; triangulin quadrate with vaguely defined depression; spermathecal duct coiled or bent between dilation and pumping region; length of sclerotized rod 1.05- 1.75.

SPECIMENS ILLUSTRATED: Genital cup: Texas: Brazos Co.:

College Station (JE); clasper: Missouri: Wayne Co. (UM0); gonocoxae: Texas: Brazos Co.: College Station (JE); spermatheca: Texas: Brazos Co.: College Station (JE).

DISTRIBUTION: Ontario, Quebec, and Maine to northern Florida; across to Arizona, Colorado (east of the Rocky

Mountains); north to North Dakota, Minnesota, Wisconsin, Michigan,

MATERIAL EXAMINED: 494 specimens from the following localities: CANADA: Ontario: Guelph (UDC); Ridgeway

(OSU). Quebec: Covey Hill (UBC). UNITED STATES:

Arizona: Coconino Co.:Oak Creek Canyon (ASU); Williams

(CU). Colorado: El Paso Co.: Rock Creek Canyon,

Colorado Springs (AMNH); Larimer Co.: Fort Collins (CSU);

Las Animas Co.: Trinidad (CSU). Connecticut: New Haven

Co.: S Meriden (CAS); Yale Bird Preserve (YLU); Tolland

Co.: Gurleyville (UCT); Mansfield Twp (UCT); Storrs (UCT).

Florida: Alachua Co.: San Felasco Hammock (UMI); Dixie

Co.: 4 mi N Old Town (UGA) ; Hwy 349 4 mi S Jct 351 (UGA); 62

Liberty Co.: Camp Torreya, T2N R7W (UMI). Georgia:

Houston Co.: Perry (UMI); Liberty Co.: St. Catherine's

Island (UGA); McIntosh Co.: Sapelo Island (UCB); Union

Co.: Levelland Mt (UGA); not located: Neel Gap (UGA).

Illinois: Champaign Co.: Urbana (INH); Cook Co.: Chicago

(CAS, INH, OSU, UMI); Glencoe (INH); Willow Springs (UMI);

Jersey Co.: Principia College, Elsah (YLU); Jackson Co.:

Carbondale (MIS); Knox Co.: Galesburg (INH); Lake Co.:

Lake Forest (CU); Mercer Co.: Keithsburg (INH); Ogle Co.:

Oregon (UMI); St. Clair Co.: New Athens (UCT); not located: Beach (INH); Chatauqua (UMI); Edgebrook (UMI);

"Ill." (ISU, UNL, WSU); "N. ill." (INH). Indiana: Brown

Co. (PDU); Clark Co. (PDU); Crawford Co.: Wyandotte (PDU);

Crawford Co. (PDU); Lake Co. (PDU); Lawrence Co.: (PDU);

Putnam Co. (PDU); Tippecanoe Co.: Monroe (PDU); Vermillion

Co. (PDU); not located: Hessville (UMI); Mineral Springs

(OSU). Iowa: Boone Co.: Ledges St Park (ISU); Johnson

Co.: Iowa City (ISU); Lee Co.: Keokuk (CU); Story Co.:

Ames (ISU, LSU, UAZ); Tama Co.: Traer (ISU, LSU); Wapello

Co.: Ottumwa (LSU, UWI); not located: Robinson (CAS).

Kansas: Atchison Co.: Effingham (AM NH); not located:

"Ks" (WSU); "Kansas" (PDU). Kentucky: Fayette Co.: Lexington (UKY); UKY Laboratory (UKY); Floyd Co.:

Prestonburg (MIS); Laurel Co.: Bald Rock (MIS); Trigg Co.

(UKY); not located: Bee Spring (AMNH); Kentucky (AMNH).

Louisiana: Jefferson Davis Parish: Jennings (CU). 63

Maine: Lincoln Co. (OHS). Massachusetts: Dukes Co.:

Martha's Vineyard (MCZ); Hampshire Co.: Amherst (CAS);

Middlesex Co.: Groton (WSU); Maiden (CSU); Suffolk Co.:

Boston (UMI); not located: Beach Bluff (CAS); Tyngsboro

(AMNH). Michigan: Alger Co.: Andrews (UMI); Onota Twp

(UMI); Allegan Co.: Swan Creek (UMI); Berrien Co.: E. K.

Warren Preserve, Sawyer Dunes (UMI); Crawford Co.: Higgins

Lake (UMI); Emmet Co. (OSU); Gladwin Co. (MIS); Huron Co.

(UMI); Ingham Co.: E Lansing (MIS); Isabella Co. (MIS);

Keweenaw Co.: Keweenaw Pt (UMI); Lenawee Co.: Clinton

(UMI); Livingston Co.: E. S. George Reserve (UMI); Manistee Co.: Portage (UKY); Midland Co. (MIS); Muskegon

Co.: Muskegon St Park (UMI); Owasippe (INH); Oakland Co.

(UAZ, UMI); Oceana Co.: Pentwater (UMI); Oscoda Co.:

T27N R2W Sec20 (UMI); Shiawassee Co. (AMNH); Washtenaw Co.:

Ann Arbor (UMI, YLU); Ann Arbor Woods (UMI); Forestry

Farm (UMI); Manchester (UMI); Washtenaw Co. (UMI); not located: "Ag. Coll. Mich." (MIS); "Mich." (UWI).

Minnesota: Mora (OSU). Missouri: Boone Co.: Ashland

Wildlife Area (UGA, UM0); Columbia (DBT, 0); Carter Co.:

Big Spring St Park (MIS); Dent Co. (UMO); Osage Co.: Elsinore (UM0); Lesterville (UM0); Linn (UMO); Randolph

Co.: 1 mi E Moberly (DBT); St. Charles Co.: Weldon

Springs (UM0); Ste. Genevieve Co. (UMO); St. Louis Co.:

Ranken (UM0); Shannon Co.: Eminence (UM0); Texas Co.

(UM0); Warren Co.: Dan Boone Nati Forest (DBT); Wayne 64

Co. (UM0). New Hampshire: Coos Co.: Mt Washington

(AMNH). New Jersey: Bergen Co.: Bear Swamp nr Ramsey

(AMNH); Middlesex Co.: Colonia (UCB); Morris Co.: Butler

(UCT); Dover (UCT); Lake Hptcong (ANNE); Ocean Co.: Beach

Haven (CAS); Holgate Unit of B. N. W. R., Long Beach Island

(INH); Ship Bottom (CAS); Sussex Co.: Lafayette (ANNE);

Union Co.: Summit (ANNE); not located: Hewitt (CAS);

NJ (AMNH, CAS); Ramapo Mts (AMNH). New Mexico: Lincoln

Co.: Ruidoso (CSU); White Mts (CSU); Santa Fe Co.:

Tesuque (AMNH). New York: Cattaraugus Co.: Gowanda

(CAS); Columbia Co.: New Baltimore (ANNE); Erie Co.:

Buffalo (CAS, INH); Colden (CAS); Lancaster (CAS); Greene

Co.: Haines Falls (CU); Jefferson Co.: Bronxville (AMNH);

Livingston Co.: Hemlock Lake (CU); Malone (CU); Nassau

Co.: Long Beach (AMNH); New York Co.: N New York and vicinity (AMNH); Ontario Co.: Honeoye (NDSU); Orange Co.:

Bear Mt (AMNH, UCR); Port Jervis (CU); Putnam Co.: West

Point (CU); Queens Co.: Queens (AMNH); Rockaway (UBC);

Suffolk Co.: Orient (CU); Sullivan Co.: Lava (CU);

Tompkins Co.: Ithaca (AMNH, CAS, CU, PDU); Ulster Co.:

Alleben (AMNH); Wayne Co.: Junius (AMNH); not located:

Nepara Pk (INH); NY (CU). North Carolina: Black Mts

(AMNH, CAS); Valley of Black Mts (ArCIE). North Dakota:

Hettinger Co. (NDSU). Ohio: Delaware Co. (OHS); Hancock

Co. (YLU); Hardin Co.: Ridgeway (ISU); Mercer Co.: Mendon

(UMI); Mercer Co. (UMI); Paulding Co.: Benton Twp (OHS); 65

Pickaway Co.: Milihof (CMNH); Richland Co.: Mansfield

(AMNH); Scioto Co. (OHS); Vinton Co.: Lake Hope St Park

(VPI). Oklahoma: Bryan Co.: Mead (OKS); Cleveland Co.:

Little River State Park (OKS); Garvin Co.: Pauls Valley

(OHS); Osage Co.: 4 mi S Hominy (OKS); Payne Co.: nr

Ripley (OKS); Stillwater (OKS); Seminole Co.: 8 mi N

Seminole (OHS). Pennsylvania: Allegheny Co.: Pittsburgh

(CMNH); Allegheny Co. (CMNH); Bedford Co.: Sulphur

Springs (AMNH); Bedford Co. (CU); Monroe Co.: Delaware Water Gap (AMNH); Washington Co. (CMNH); Westmoreland Co.:

Jeannette (CSU); not located: Penn. (AMNH, TAM). South

Carolina: Greenville Co. (JE). South Dakota: Brookings

Co.: White (ISU). Tennessee: Knox Co.: Knoxville (ISU);

Shelby Co.: Shelby Forest St Pk (MIS); Stewart Co.: Land

Between the Lakes, nr Model (MIS). Texas: Brazos Co.:

College Station (JE, TAM); Dallas Co.: Dallas (AMNH);

Galveston Co.: Galveston (CAS, ISU); Gillespie Co. (OHS);

Lubbock Co.: Lubbock (TTU); Mills Co.: E of Goldthwaite

(JE); Montgomery Co.: The Woodlands (DET); San Saba Co.:

Gormon Falls (JE); not located: "Tex." (INH, MCZ).

Virginia: Bland/Wythe Co.: Summit Walker Mt 3950' (=I).

West Virginia: Pocahontas/Randolph Co.: Cheat :'Its (CMNH).

Wisconsin: Bayfield Co. (UWI); Clark Co. (UWI); Dane Co.:

Madison (UWI); Dane Co. (UWI); Greenlake Co. (UWI);

Lakewood Co. (MIS); Polk Co.: Gibson Lake (UWI); Shawano

Co. (MIS); Waukesha Co. (UWI); vTaupaca Co. (UWI) ; Wood 66 Co.: Nepeo Lake (UWI); S Nepeo Rd (UWI); not located: "wis.0(wsu).

This species is rather constant morphologicallyover its range. Some variation in the number and intensity of

the metallic green punctureson the pronotum exists. The

dorsal arm of the clasper also varies in itsorientation with respect to the ventralarm. The former is parallel

to the latter in some specimens, while in othersthe dorsal arm points dorsolaterad. Female external genitalia are very uniform, but minor differences in the spermathecal

bulb and coiling of the spermathecal duct exist.

A. cynicus' distribution overlaps with A. bracteatus'

in the northeastern and north centralU. S., and eastern

Canada. Most of this area falls into a suture zone,or area where hybridization of related species occurs due to the removal of a barrier, believed in thiscase to be a continuous forest (Remington 1968). As detailed habitat and phenological studies of these species have not been made, the occurrence of hybridizationor its potential is not known; however, among 950 specimens, I encountered none which appeared to be intermediates.

The eggs and nymphs of this specieswere described by

Whitmarsh (1916) and Jones andCoppel (1963). Results of life history studies are shown in Table 1 andprey records are listed in Table 2. Stoner (1917) observed this species in woodlands or in small clearings in the OkobojiLakes 67 area of northwest Iowa.

GENUS PODISUS HERRICH - SCHAEFFER

Podisus Herrich - Schaeffer 1853: 296. Type species

Podisus vittipennis H.- S. 1853 (synonym of Arma

nigrisnina Dallas 1851); by subsequent desianation

(Kirkaldy 1909: 13).

Telepta Stal 1860: 10. Type species = Telepta crassimarao Stal 1860; by subsequent designation (Kirkaldy 1909:

18).

Podisus, subgenus Podisus Stal 1870: 49; Van Duzee 1917:

78.

Podisus, subgenus Tylospilus Stal 1870: 52; Van Duzee 1917:

81. Type species = Arma chilensis Spinola 1852; by

subsequent designation (Schouteden 1907: 70, as

Apateticus chilensis).

Apateticus, subgenus EulDodisus Schouteden 1907: 71. Type

species = Arma modesty Dallas 1851 (synonym of Pentatoma

maculiventris Say 1831); Schouteden 1907: 71, as

Apateticus modestus.

Apateticus, subgenus Tylospilus Schouteden 1907: 73;

Kirkaldy 1909: 21. Apateticus, subgenus Podisus Kirkaldy 1909: 18. 68

DIAGNOSIS: Generally less than 12 mm long; juga parallel, length equal to tylus; males lacking rugulose setoseareas on sternites IV - VI; fore femora unarmed; fore tibiae not dilated. Claspers generally triangular, theca subequal to thecal shield; conjunctival appendages entirely membranous.

Female triangulin covered by first gonocoxae.

DESCRIPTION: Body length: male 7.0 - 13.5; female 8.0 -

14.0. Humeral width: male 4.1 - 7.5; female 4.4 - 8.3.

Shape ovate, most species widest at humeri. Dorsum yellow to brown, punctation rufous, brown or fuscous; venter pale yellow to dark brown, impunctate or punctation rufous to brown. Punctation coarser dorsally than ventrally. Head moderately declivous, dorsally planar, subtruncate to rounded apically, length and width subequal; tylus and juga of equal length. Rostrum slender to moderately stout, extending to or between metacoxae; second segment longest, fourth segment darkest. Antennae slender, pale yellow to light brown or rufous, setulose, subequal to half body length; second segment usually longest; remainder subequal. Pronotum declivous forward of humeri; anterolateral margins concave-dentate to straight. Humeri blunt to spinose; spines notched or entire, directed laterad or cephalad, elevated or level.

Scutellum longer than wide; fovea in each basal angle; frena reaching or surpassing middle; at times with medial irregular slightly raised pale longitudinal line. 69

Hemelytra usually rufous ventrally at apex and on embolium.

Membrane clear to pale yellow or brown, with or without darker median longitudinal band. Legs concolorous with venter, at times rufous, finely punctate; fore femora lacking spines or tubercles; fore tibiae not dilated. Venter with or without variable series of small black dots laterally and/or mesally; often sternite VI with large median black spot; males lacking rugulose setose patches on sternites IV - VI. Ventral spine present, directed forward, variable size. Duct of metathoracic scent gland short, straight or curved forward in apical half. Genitalia: male genital cup shallow, variable in shape, contents visible or obscured in caudal view; clasper with short shaft attached to cup wall, curved dorsad or laterad into flat, expanded area of variable shape; median penial lobes disk-like; 1 pair membranous conjunctival appendages. Female first gonocoxae quadrate; spermatheca with wide dilation containing sclerotized rod enclosing duct leading into pumping region; bulb variable in shape; flanges small to prominent.

DISTRIBUTION: Confined to the Nearctic and Neotropical regions. Nine species are included in the Nearctic fauna. P. placidus, P. maculiventris, P. brevispinus, P. fretus, and P. serieventris are exclusively NearCtic. sacitta,

P. concrrex, and P. acutissimus are predominantly

Neotropical but extend into the Nearctic reaion. 70

mucronatus is known only from Florida and Cuba.

Key to the Nearctic Species of Podisus

1. Pronotum with red to fuscous transverse band; base of

scutellum with 2 - 3 distinct ivory or rufous levigate

spots; male genital cup tilted forward, obscuring

contents; ventral border of cup subtended by median U-

or V- shaped notch (fig. 7b) . (subgenus Tylospilus)... acutissimus, p. 72

- Pronotum without transverse band; scutellum lacking

distinct basal spots....(subgenus Podisus) 2

2. Humeral spines curved forward; anterolateral pronotal

margins forward of spines explanate, ivory, dentate

(fig. 6a); apex of scutellum ivory; found in Florida,

Cuba mucronatus, p. 83

- Humeral spines, if present, directed laterally, at most curved forward only slightly; tip of scutellum

not ivory 3

3. Anterolateral pronotal margins straight, humeri blunt

(fig. 13); membrane of hemelytra lacking dark

longitudinal band nlacidus, p. 142

Anterolateral pronotal margins concave - dentate;

humeri variable 4

4. Ventral spine short (fig. 9e, 11 ) 5

Ventral spine extending between hind coxae ig. 8f) 6 71

5. Color mottled brown - aray; humeri bluntly rounded

(fig. 9a); clasper with subacute dorsal and lateral

apices (fig. 9c) serieventris, p. 114

Color lighter brown than preceding, not mottled;

humeri narrowly rounded (fig. 11a); clasper with

rounded dorsal and lateral apices (fig. 11a)

brevistinus, p. 125

6. Humeri bluntly rounded (fig. 12a); venter with

irregular reddish brown patches; clasper with broadly

trianaular dorsal portion (fig. 12e).. fretus, p.138 Humeri subacute to spinose; venter without reddish

patches 7

7. Humeral spines notched apically (fig. 14a)

sagitta, p. 152

- Humeri subacute or with spines entire 8

8. Humeral spines fuscous or rufous at tips; directed

laterad, slightly elevated; anterolateral pronotal

margins levigate, swollen, dentate (fig. 10a); found

in :lexico, Central America, and northern South

America conarex, p. 122

Humeri subacute to spinose; spines when present may be

broadly curved forward or directed laterad (fig. 8a);

widely distributed in the U. S. and Canada maculiventris, p. 87 72

Subgenus Tylospilus Stal

Podisus, subgenus Tvlosnilus Stal 1870: 52; Van Duzee

1917: 81. Type species = Arma chilensis Spinola 1852;

by subsequent designation (Schouteden 1907: 70, as Anateticus chilensis).

Aoateticus, subgenus Tvlosnilus Schouteden 1907: 73;

Kirkaldy 1909: 21.

DIAGNOSIS: Base of scutellum with 2 - 3 distinct, glabrous rufous spots; frena reaching mid-scutellum. Genital cup tilted forward; with U- or V-shaped median notch subtending ventral border.

Podisus acutissimus Stal (Figs. 7, 15i, k; Map 7)

Podisus (Tvlosnilus) acutissimus Stal 1870: 53; Distant

1880: 40, pl. 2, fig. 22: Van Duzee 1917: 81.

Podisus acutissimus Kirkland 1898: 435; Van Duzee 1904: 72.

Tvlosnilus acutissimus Uhler 1276: 283.

Apateticus (Tvlosnilus) acutissimus Schouteden 1907: 73;

Kirkaldy 1909: 21.

Podisus rubromaculatus Distant 1880: 41, pl. 7, fig. 1

NEW SYNONYI:Y.

TYPES AND TYPE LOCALITIES: Podisus acutissimus Seal, 2 syntypes (Riksmuseet); female, fair condition: "Mexico /

Salle. / Type / Allotypus (red) / 408; 82 (red) / 73

Riksmuseum; Stockholm (green)." Male, fair condition:

"Mexico / Sall6 / Type / Paratypus (red) / 409; 82 (red) /

Riksmuseum; Stockholm (green)." The type locality is

Mexico. The red labels are museum labels with no nomenclatural significance; however, the specimens are listed as syntypes on themuseum invoice (P. Lindskog, pers. comm.). I will propose that the male be given lectotype status in a subsequent publication. Podisus rubromaculatus Distant, type, male, (Naturhistorisches Museum, Vienna), very good condition: "Delimek, Mexico; 1871 / rubromaculatus, det. Distant / rubromaculata / Coil. Nat. Mus. Wien."

DIAGNOSIS: Red to fuscous transverse band on pronotum; base of scutelluM with 2- 3 glabrous ivory or rufous spots; red, fuscous, and ivory patterns on apical half of scutellum. Claspers oriented so flat, broadly sickle- shaped surfaces facing medially.

DESCRIPTION: Body length: male 7.0 - 9.5; female 8.0 -

10.5. Humeral width: male 4.3 - 5.7; female 5.0 - 6.6. Dorsum pale yellow to light brown with distinctive red, ivory, and fuscous markings; punctation very fine, overall appearance smooth; venter concolorous or lighter than dorsum. Head rounded, slightly tapered apically; width: male: 1.5 - 2.0; female 1.7 - 2.1; length: male 1.4 -

1.8; female 1.6 - 2.0. Antennae yellow to brown; seument 74

II sometimes fuscous to blaCk. Pronotum with red to black transverse band extending onto humeri; band varies in width and color intensity; anterolateral margins of pronotum denticulate, straight. Humeri produced into acute, polished spines which may curve forward. Scutellum: width: male 1.9 - 2.6; female 2.3 - 3.0; length: male

2.3 - 3.2; female 2.7 - 3.6; basally with 2 - 3 glabrous ivory or red spots; median longitudinal pale line; apical

1/3 fuscous to black on either side of pale line with pinkish area laterad and distad of black. Hemelytra with small black dot in apical 2/3; membrane hyaline with dark band. Venter with small black spots laterally; ivory or red spots may be present on pleura. Duct of metathoracic scent gland short, straight. Legs concolorous or darker than venter; tarsi darkest; tibiae smooth dorsally.

Ventral spine stout, extending to anterior margins of hind coxae. Male genitalia: width of genital cup 0.96 - 1.22; caudal opening small; dorsal face narrow; sinuous ventral border setose; claspers pointing dorsad, shaped like broad, blunt sickles with surfaces directed medially; genital plates subquadrate, mesal margins toothed, irregular; dorsal margins deflexed with tooth at dorsolateral corner. Female genitalia: gonocoxae subquadrate, width 0.42 - 0.53; length 0.40 - 0.53; spermatheca small; bulb loear-shaped to oval with prominent flanges; length sclerotized rod 0.33 - 0.68. 75

SPECIMENS ILLUSTRATED: Humerus, duct of scent gland:

Arizona: Cochise Co.: 5 mi H Portal 5400' (ASU); genital cup: Arizona: Cochise Co.: 5 mi H Portal 5400' (AMNH); genital plate, clasper, dorsal view of genital cup:

Arizona: Cochise Co.: Painted Canyon Ranch, Chiricahua

Mts (AMNH); spermatheca: Arizona: Cochise Co.: Painted

Canyon Ranch, Chiricahua Mts (AMNH).

DISTRIBUTION: Southern California, Nevada, Arizona, New

Mexico, Texas, Oklahoma, Arkansas, Kansas, Missouri,

Florida, Mexico, Costa Rica, Curacao, Dominican Republic,

Bahamas (Map 7).

MATERIAL EXAMINED: 772 specimens from the following localities: BAHAMAS: Andros Island: San Andros (LYE).

COSTA RICA: Guanacaste: 3 km NW Canas (TAM) . CURAgAO:

Pisadera Bay (AMNH). DOMINICAN REPUBLIC: Santiago:

Santiago (WSU). MEXICO: Aguascalientes: 4 mi SW

Aguascalientes (AMNH); Pabelln (UCB). Bala California:

Chapala, dry lake (CAS); 14 mi S El Arco Mine (CAS); 15 mi

N El Refugio (CAS); Las Arrastras (ASU); 10 mi E Mesguital

(CAS); 10 mi S Punta Prieta (CAS); San Domingo (CAS); 15 mi

N San Ignacio (CAS); Venancio (CAS). Chiapas: Tuxtla-

Guti4rrez (AMNH). Chihuahua: Catarinas (AMNH); 239 km S

Ciudad JuArez (ANNH); Moctezuma (UCB); Salaices (ANNH);

Santa Clara (AMNH); Santa Clara Canyon, 5 mi H Parrita

(UCB); 10 mi S Villa Ahumada (UCB). Coahuila: 5 mi NE 76

La Rosa (MIS); Paila (AMNH); 4 mi W Saltillo (TTU).

Durango: 5 mi NE Donato Guerro (TTU); 5 mi H Guadalupe

Victoria (TAM); 3 mi SE Nombre de Dios (TAM); Pedricena

(AMNH); 5 mi S Pedricena (TAM); Tapias (MIS); Tiahualillo

(ISU, TAM). Guanajuato: San Miguel Allende (AMNH).

Guerrero: 49 km N Chilpancingo (DBT); Rio Balsas (CAS).

Hidalgo: 5 mi E Jacala (CAS); 7 mi S Jacala (TAM).

Jalisco: Ajijic (UCB); 16 km N Autlan (TAM); 37 mi S

Guadalajara (UCB); 7 mi S Mazamitla (CAS); SE slope Mt

Colima (CAS);1 mi NE Villa Hidalgo (AMNH); 22 km NE

Zapotlanejo. Morelos: 12 mi E Cuernavaca (KU); 5 mi S

Cuernavaca (CU); Puente de Ixtla (CAS). Nayarit: 20 mi S Acaponeta (TAM); 26 mi S Acaponeta (UCR); San Bias (CAS);

Tepic (CAS, MIS); 4 mi NW Tepic (ANNE); 17 mi NW Tepic

(CAS). Nuevo LeOn: 9 mi H Iturbide (TAM); 15 mi W

Linares (TAM); 23.6 mi SW Linares (TAM); 7.5 mi S Monterrey

(JE, TAM); 41 mi S Saltillo (KU); 10 mi S Jct 57 & 60

(TTU). Oaxaca: 3.2 mi S Ocotlan (TAM); 10.3 mi S Rio

Grande (Hwy 131) (JE). Puebla: 25 mi SE Puebla (CAS);

6 mi SW TehuacSn (TAM). Queretaro: Queretaro (LCD).

San Luis Potosi: 40 - 50 mi NW Ciudad del Maiz (CAS); El Salto (KU); 6 mi E Matehuala (UCB);1 mi S San Lorenzo

(TAM); 17 mi NE San Luis Potosi (OSU); 40 mi S San Luis

Potosi 5700' (OSU). Sinaloa: Mazatlgn (CAS). Sonora:

Minas Nuevas (AMNI-1); 15 mi E Navojoa (LACK); Saric (CAS).

Tamaulipas: Bocatoma, 6 mi S Gomez Farias (DET); 77

28 mi S Ciudad Victoria (CAS); Gomez Farias (DBT); 25.4 mi

SW Jaumave (TAM); 8 mi E Padilla, Rancho Santa Ana (UMI).

Veracruz: Ciudad Mendoza (AMNH); Fortin (AMNH); Minatitlgn

(MIS); Orizaba (ISU); Puente Nacional (UGA). Yucatan: Cordeleria Mayaan (AMNH); Dolores Otero (AMNH); Tixkokob

(AMNH). Zacatecas: Fresnillo (AMNH); 9 mi S Fresnillo

(UCB); Sain Alto (AMNH). UNITED STATES: Arizona: Coconi- no Co.: Gray Mt (AMNH); Oak Creek Canyon (KU); Cochise Co.:

Douglas (AMNH, CSU, KU); 8 mi E Douglas (UAZ); 10 mi E

Douglas (CAS); Elfrida (SDCM); Painted Canyon Ranch,

Chiricahua Mts (AMNH); 10 mi E Portal (ASU);1 mi N Portal

(UGA);1 mi NE Portal (UCR); 2 mi NE Portal (ASU); 1 mi SW

Portal (UCB);1 mi W Portal (UCB); 3 mi W Portal (OSU);

Portal (AMNH, UCB, UCR); 5 mi W Portal, SWRS (AMNH, ASU,

LACM, OSU, UGA, UID); San Bernardino Ranch (KU); 15 mi E

Sonorita (ASU); Texas Canyon, 11 mi W Apache (UCR); Tomb- stone (KU); Wilcox (UAZ); Benson (CAS, KU); Carr Canyon, Huachuca Mts (AMNH); Gila Co.: Globe (KU); Graham Co.:

Bonita (UAZ); Dripping Spring, Whitlock Mts (UAZ); Safford

(ASU); San Carlos Lake (AMNH); Maricopa Co.: Buckeye (ASU); Mesa (ASU); Tempe (ASU); Wickenburg (OHS, UCT); Mohave Co.:

Hackberry Wash (OSU); Hualapai Mt Park (UID); 23 mi E

Kingman (KU); 27 mi SE Moyucca (UCB); Pima Co.: 15 mi S Ajo (LACM); Browns Canyon, Baboquivari Mts (AMNH); Lower

Madera Canyon, Santa Rita Mts (UID); Madera Canyon, Santa

Rita Mts (AMNH); Sabino Canyon, Santa Catalina Mts 78

(CAS, OSU, UAZ); Santa Rita Mts, Rae Res (UAZ); San Vicente

(AMNH); Tucson (AMNH, ISU, OHS, OSU); Tucson, Bear Canyon

(CAS); 15 mi E Tucson (UCB); 28 mi N Tucson (UCB); nr Vail

(AMNH); Pinal Co.: Antelope Pass (ASU); Elroy (UAZ);

Santa Cruz Co.: Nogales (CAS, ISU); Patagonia (CAS, UAZ);

Yavapai Co.: Congress Jct (OHS); Prescott (ASU, OHS); 10 mi N Prescott (AMNH); Yuma Co.: San Luis (UID); not lo- cated: "Arizona" (CU); B.T. Arboretum (KU); Carrizo (UAZ);

Charleston (CAS); Christmas (AMNH); Cortaro (UAZ); 22 mi S

Mt View (UAZ); Pinal Mts (ISU); Stewart Lake (UAZ).

Arkansas: Crittenden Co.: Turrell (LSU). California:

Imperial Co.: Bard (UCB); El Centro (UCR); Potholes (CAS);

Yuma Valley (UMI); Inyo Co.: China Ranch (LACM); Kern Co.:

16 mi S Weldon (UCB); Riverside Co.: Palms to Pines Hwy

(UCB); San Bernardino Co.: 10 mi N Earp (UCB); Needles

(CAS); San Diego Co. (SDCM). Florida: F. Capron (INH).

Kansas: Ford Co.: Dodge City (KU). Missouri: Boone Co.:

Columbia (UMO). Nevada: Lincoln Co.: Alamo (OSU).

New Mexico: Catron Co.: Apache Creek (OSU); Chaves Co.:

Mescalero Sands (AMNH); Roswell (KU); Doria Ana Co.: Las

Cruces (AMNH); 11 mi S Road Forks (UMO); Hidalgo Co.:

Granite Gap, Peloncillo Mts (ASU); Lordsburg (AMNH); 8 mi

SE Rodeo (AMNH); Lincoln Co.: 20 mi NW Carrizozo (AMNH);

Ruidoso (MIS); Torrance Co. (KU). Oklahoma: Jackson Co.:

1 mi S Altus (OKS); Lincoln Co.: Sparks (OKS); 79

McIntosh Co.: Eufaula (OHS); Payne Co.: Stillwater (OKS);

Texas Co.: Goodwell (OKS); Texas Co. (OKS); Woodward Co.:

Fort Supply, Beaver River (OKS). Texas: Blanco Co.: 8 mi

NW Johnson City (JE); Brazoria Co. (KU); Brazos Co.:

College Station (ISU, JE, LSU, TAM, UMI); Brazos Co. (MIS);

Brewster Co.: Alpine (AMNH, KU, LSU); Big Bend (KU);

Big Bend Nati Park, Chisos Basin (TTU, UID); Big Bend Nati

Park, Pine Canyon 5100' (TTU); Big Bend Natl Park (TAM);

Chisos Mts (OHS, UAZ, UCT, UID); Chisos Mts, Big Bend Natl

Park (AMNH, CAS, OSU); Chisos Mts, Big Bend (CAS); Green Gulch, Big Bend Natl Park (TAM, TTU); Marathon (AMNH); 9 mi S Marathon (AMNH); 14 mi E Panther Jot (LACM); Tenapin

Tanks, Big Bend Natl Park (AMNH); The Basin, Big Bend Natl

Park (AMNH, TAM); Brooks Co.: Falfurrias (UCB); Burleson

Co. (TAM); Burnet Co.: Ink's Lake State Park (UMO);

Cameron Co.: Brownsville (AMNH, CAS, CMNH, CSU, CU, INH

KU, OKS, UCB, UCT, UMI); 6 mi E Brownsville (TAM); Langtry

(LSU); Port Isabel (CAS); Sabal Palm Grove Sanct (DBT, UGA);

White Wing Reserve, Brownsville (TAM); Cameron Co. (KU);

Cochran Co. (TTU); Collin Co.: McKinney (TAM); Dickens Co.: 7 Bar Ranch, Dickens (TTU); Dimmit Co. (TAM); El Paso Co.:

Fabens (LSU); Erath Co.: Stephenville (TAM); Floyd Co.:

12 mi NE Floydada (TTU); Floyd Co. (MIS); Gaines Co. (TAM);

Gillespie Co.: Fredericksburg (JE); Hidalgo Co.: Bentsen Rio Grande Valley State Park (DBT); 12 mi W Mission (KU);

On Rio Grande, Hidalgo (UAL); Santa Ana Wildlife Refuge 80

(TAM); Weslaco (TAM, UMI); Hidalgo Co. (KU); Jeff Davis

Co.: Davis Mts (KU, OHS, UCB); Fort Davis (LSU, UID);

6 mi SE Fort Davis (TAM); 10 mi SE Fort Davis (TAM); Limpia

Canyon, Davis Mts (LACM); 22 mi S Van Horn (OSU); Kerr Co.:

Kerrville (TAM); Kleberg Co.: Kingsville (CU); Lubbock

Co: Lubbock (TTU); Lubbock Co. (TTU); McLennan Co.:

Mart (MIS); Waco (AMNH, TAM); Menard Co. (KU); Nueces Co.:

Corpus Christi (TAM); Pecos Co.: Fort Stockton (OHS);

Presidio Co.: Alamito Creek, 5 mi SE Presidio (TAM); Blata

(TAM); Chisati Mt (UMN); Presidio (OKS, TAM); 3 mi N Presi- d1D(JE, TAM); 13 mi N Presidio (TAM); 2 mi SE Presidio (TAM);

Shafter (TAM); Presidio Co. (KU); Reeves Co.: Pecos (OKS); TAM Res Stn at Pecos (TAM); Robertson Co. (TAM); San

Patricio Co.: 4 mi SW Mathis (TAM); Sinton (TAM); Welder

Wildlife Refuge (TAM); Starr Co.: Falcon State Park (TAM);

Sutton Co. (KU); Swisher Co. (TTU); Tarrant Co.: Fort

Worth (CAS); Terrell Co.: Sanderson (KU); Travis Co.:

Austin (CAS, WSU); Upshur Co. (TAM); Uvalde Co.: Sabinal

(TTU); Uvalde (CAS); Val Verde Co.: Del Rio (KU); Val

Verde Co. (UCT); Victoria Co. (KU); Walker Co.: Ellis

Prison Unit (JE); Webb Co.: Laredo (UCT); 30 mi NW Laredo

(WSU); not located: Corrizo Springs (KU); Dryden (CAS);

Langtry (LSU); "Texas" (CAS, INH); Valentine (KU). 81 The color and size of the pronotal band and markings

on the scutellum are variable. The former ranges from very

wide and dark, sometimes with the margins lighter,to thin

and red, often in specimens from thesame series.

Specimens with a thin, red pronotal bandgenerally have

red, rather than ivory, spots basallyon the scutellum.

Distant's type of P. rubromaculatus isan example. The

humeral spines may be directedlaterally or curved forward. Some individuals from southern Mexico, Central

America, and the Caribbeanare paler than northern

representatives. Barber and Bruner (1932) reported that

Uhler misidentified some light specimens ofP. acutissimus

taken in St. George, Grenada, West Indiesas P. aaumeri Distant. I have not seen these insects.

Male genitalia are constant throughout therange. Females show minor variations in the shape of the spermathecal bulb.

Gillette and Baker (1895) list this speciesfrom Colorado. No specific locality was aiven and I examined no Colorado specimens. As P. acutissimus occurs in Kansas and New Mexico, it probably alsoranges into southern Colorado. I have not indicated its occurrence in Colorado on the distribution map (Map 7).

Stoner et al. (1974b) reared this species at 5 temperatures and found differences in stadial duration and survivorship. Results for the optimum selected temperature 82

(26.1°C) are given in Table 1. In a related study, Stoner et al. (1974a) found that whereas P. acutissimus could not survive to adulthood solely on plant food, nymphs successfully reached the second instar.

Ruckes (1938) reported that P. acutissimus is uncommon and of local occurrence in New Mexico. His specimens were collected on Holcus halepensis L.

(Gramineae).

Small numbers of this species have been collected from alfalfa, barley, cotton, and sorghum in Arizona

(Butler and Werner 1960; Stoner et al. 1974a, b). Because of its scarcity, this species is not considered a major predator in Arizona crops (Butler and Werner 1960).

Virtually nothing is known of its habits, thus its value as a component of augmentation and conservation schemes for biological control should not be dismissed without further study.

Subgenus Podisus Herrich - Schaeffer

DIAGNOSIS: Coloration various shades of brown; frena exceeding mid-scutellum; humeri variable. Claspers generally triangular. 83

Podisus mucronatus Uhler

(Fig. 6; Map 6)

Podisus mucronatus Uhler 1897a: 386; Kirkland 1898: 437.

Apateticus (Eupodisus) mucronatus Schouteden 1907: 72.

Apateticus (Podisus) mucronatus Kirkaldy 1909: 19.

TYPE AND TYPE LOCALITY: Podisus mucronatus Uhler, syntype,

(no type label), male, good condition (USNM): "Cuba /

P. R. Uhler Collection / 381 / Podisus mucronatus Uhl.;

Cuba; det. Uhler / Podisus mucronatus; Cuba Uhler." I will propose that this specimen be given lectotype status in a future publication. Uhler notes in his description that he examined upwards of a dozen specimens collected in Cuba by him and John Gundlach, and in southern Florida by friends.

DIAGNOSIS: Humeral spines prominent, directed forward; apex of scutellum ivory; punctation coarse; anterolateral margins of pronotum explanate, ivory, toothed.

DESCRIPTION: Body length: male 8.5 - 11.0; female 9.5 -

12.0. Humeral width: male 5.1 - 6.2; female 5.2 - 6.4.

Dorsum yellowish brown or paler with brown to reddiSh punctation; Venter pale yellow, coarsely and concolorously punctate. Head with borders anterior to eyes narrowly black; edges of tylus dark brown to black, concolorous with jugal punctation; tylus and juga pale, 84 the latter to inner posterior margins of eyes; width: male 1.7 - 2.0; female 1.8 - 2.1; length: male 1.6 - 1.8; female 1.5 - 1.8. Antennae pale yellow except darker apical segment. Pronotum coarsely punctate; glabrous ivory spot posterior to each callus, joining median pale band extending between and anterior to calli; margins forward of humeri explanate, ivory, toothed.Humeri produced into prominent forwardly curved fuscous spines.

Scutellum: width: male 2.5 - 3.0; female 2.6 - 3.2; length: male 3.0 - 3.6; female 3.1 - 4.0; basal, lateral ivory-colored spots at times coalescent. Legs concolorous with venter; tarsi darker.Ventral spine extending between hind coxae. Male genitalia: width of genital cup

1.08 - 1.22; lateral lobes reduced; ventral area turned abruptly dorsad creating broad, slightly concave region below setose, sinuate ventral border. Proctiger with C- shaped ridge dorsally curving mesally. Claspers V-shaped, directed dorsolaterally; mesal arm of "V" slender, acute; lateral arm wider with pointed apical process. Genital plates subquadrate except for narrow lateroventral extension; mesal margin variously reflexed; surface of plate with irregular scalloping and/or flattened tubercles.

Female genitalia: first gonocoxae rugose; margined mesally with dark brown; width 0.53 - 0.64; length 0.48 -

0.53; bulb oval - elongate with small flanges; length of scierotized rod 0.78 - 1.02. 85

SPECIMENS ILLUSTRATED: Humerus, clasper: Florida:

Broward Co.: Hallendale (AMNH); genital cup: Florida:

Collier Co.: Naples (AMNH); proctiger: Florida: Broward

Co.: Hallendale (AMNH); gonocoxae: Florida: Orange Co.:

Winter Park (AMNH); spermatheca: Florida: Broward Co.: Hallendale (AMNH).

DISTRIBUTION: A Caribbean species collected in Florida and Cuba (Map 6).

MATERIAL EXAMINED: 108 specimens from the following localities: CUBA: Baragua (AMNH); Santiago de las Vegas

(UCB); Trinidad, Las Vegas (AMNH). UNITED STATES:

Florida: Alachua Co.: Gainesville (PSU); Brevard Co.:

1.5 mi N Eau Gallie (UMI); Broward Co.: Hallendale (AMNH);

Charlotte Co.: Charlotte Harbor (AMNH); Charlotte Harbor

Area (UMO); Collier Co.: Naples (AMNH); Dade Co.:

Biscayne Bay (AMNH, CAS, OHS); Miami (PSU); Paradise Key

(UMI); Hendry Co.: La Belle (KU); Highlands Co.: Archibold

Biological Stn (CAS); Indian River Co.: Indian River

(AMNH); Sebastian (AMNH); Lee Co.: Fort Myers (CAS, PDU);

Manatee Co.: Cortez Beach (UMI); Monroe Co.: Upper

Matecumbe Key (DBT); Okeechobee Co.: Okeechobee (AMNH);

Orange Co.: Winter Park (AMNH); Osceola Co.: Kissimee

(PDU); Palm Beach Co.: Jupiter (AMNH); Pahokee (CU, UCB);

Palm Beach (CAS); Royal Palm Park (OSU, PDU); Pinellas Co.:

Dunedin (CAS, PDU, UNN); Polk Co.: 0.75 mi E Hesperides 86

(UMI); Lake Hamilton (UMI); Seminole Co.: Oveida (KU);

Sanford (ISU, KU, UAZ); Volusia Co.: Ormond (PDU); not located: Brighton (CU); Cape Sable (PDU); Coconut Grove

(AMNH); Coxam (PDU); Goon Swamp (PSU); Lake Worth (AMNH); Princeton (UCT).

This species is rather constant morphologically over its aeographic range. Minor differences exist in the degree to which the humeral spines point forward and in the red coloration of the dorsal punctures. Male genitalia are also uniform. Females show small differences in the shape of the spermathecal bulb.

Little is known of this species' life cycle or habits.

Blatchley (1926) collected single specimens of P. mucronatus on cabbage palmetto and foliage at the edges of sandy fields in Florida. Hayslip et al. (1953) recorded it as a predator of caterpillars attacking crucifers. No particular species was mentioned as prey of P. mucronatus, but the authors list Spodoptera fructiperda (J. E. Smith)

(fall armyworm), Trichoplusia ni (Hubner) (cabbage looper),

Agrotis ipsilon (Hufnagel) (black cutworm), Feltia subterranea (F.) (granulate cutworm), and Hellula rogatalis (Huist) (cabbage webworm) as lepidopterous pests of crucifers.

Genung (1960) reported P. mucronatus and P. maculiventris as the most important insect predators of

Autoplusia eaena (Guen4e) (bean leafskeletonizer). A 87 cross-mating pair of P. mucronatus and P. maculiventris was seen feeding on caterpillars of

(Grote) on Ficus in Florida (Genung 1959). Maddox et al.

(1971) found the same 2 species preying on Agasicles hygrophila Selman & Vogt, a chrysomelid imported to control alligatorweed in Florida waterways. As only small numbers of Podisus were found, the extent of herbivore population reduction was not known.

Podisus maculiventris (Say)

(Figs. 8, 15b, d, h, j, Map 8)

Pentatoma maculiventris Say 1831: 11; reprint 1899: 307.

Arma spinosa Dallas 1851: 98.

Arma modesta Dallas 1851: 101. NEW SYNONYMY.

Arma pallens Stil 1859: 222.

Podisus maculiventris Uhler 1899: 307. Apateticus (Eupodisus) maculiventris Schouteden 1907: 72.

Apateticus (Podisus) maculiventris Kirkaldy 1909: 19.

TYPES AND TYPE LOCALITIES: Say's specimen(s)is destroyed. The type locality is Louisiana. Arma spinosa Dallas, type, female, poor condition (BM): "Type (circled) / N. America;

63-90 / 4. Arma spinosa." Arma modesta Dallas, type, male, poor condition (BM): "Type (circled) / 5. Arma modesta

/ 698 / small green diamond-shaped label with "R" written on reverse." Type localities listed by Dallas are Trenton 88

Falls, New York, and Cincinnati, Ohio. Arma pallens St&l,

syntype, female, good condition (Riksmuseet): "California / Kinb. / Type / Paratypus (red) / 413;82 (red) /

Riksmuseum, Stockholm (green)." The type locality is San

Francisco, California. I will propose that the A. pallens

syntype be given lectotype status, as the red type labels are for museum use and of no nomenclatural significance

(P. Lindskog, pers. comm.). I also will propose a neotype from Say's type locality (Louisiana).

DIAGNOSIS: Color brown to gray, not mottled, humeri subacute to spinose, may be curved forward; ventral spine long, extending between hind coxae. Claspers triangular, concave laterally.

DESCRIPTION: Body length: male 9.5 - 12.5; female 10.5 -

14.0. Humeral width: male 5.2 - 7.0; female 5.5 - 8.0.

Dorsum ochraceous to grayish brown with darker punctation; venter ivory to light yellow; punctation red to brown.

Head: width: male 1.8 - 2.5; female 2.1 - 2.5; length: male 1.6 - 2.2; female 1.8 - 2.3. Antennae rufous to piceous; incisures paler; segment II usually palest.

Pronotum with concave, dentate anterolateral margins.

Humeri acute to spinose. Scutellum: width: male 2.4

3.3; female 2.9 - 3.6; length: male 3.2 - 4.3; female

3.4 - 5.0. Membrane with dark band. Legs concolorous with venter or darker; sometimes reddish. Venter with row 89 of black spots laterally, mesally, variable in size; females usually with large black mesal spot on sternite

VI. Ventral spine extending between hind coxae.Male genitalia: width of genital cup 1.26 - 1.58; lateral lobes small, rounded; apically deflexed. Claspers triangular, laterally concave, shallowly or with point of inflection midway along serrulate lateral margin; dorsal and lateral apices subacute to acute. Genital plates quadrate, indented laterally; surface ridged, tuberculate. Female genitalia: width of first gonocoxa

0.53 - 0.80; length 0.53 - 0.58. Bulb elongate, often curved or slightly constricted; flanges prominent. Length of sclerotized rod 0.76 - 1.10.

SPECIMENS ILLUSTRATED: Spinose humerus: Louisiana: Baton

Rouge Parish: Baton Rouge (LSU); subacute humerus:

Oklahoma: Latimer Co.: Wilburton (OKS); head and duct of scent gland: Louisiana: Baton Rouge Parish: Baton Rouge

(LSU); rostrum: Louisiana: Cameron Parish: Franklin

(LSU); ventral spine, genital cup: Louisiana: Baton

Rouge Parish: Baton Rouge (LSU); clasper, genital plate, median penial lobes: Oklahoma: Garwin Co.: Strang (OKS); spermatheca: Louisiana: Baton Rouge Parish: Baton Rouge

(LSU).

DISTRIBUTION: Widespread throughout the United States and

Canada (Map 8). 90

MATERIAL EXAMINED: 4071 specimens from the following selected localities: CANADA: British Columbia: Saanich

District (UBC); Victoria (INH). Manitoba: Manitoba (UAZ).

Nova Scotia: Boisdale (AMNH); Kentville (CAS); Kings Co.

(UBC). Ontario: Byron Bog, London (UAL); Guelph (UBC);

Ridgeway (ISU); Rondeau Park (UMI); Thessolon Lake Short

(AMNH); Toronto (ISU). Quebec: Charlevoix Co. (UBC);

Montreal (CMNH, JE, KU, UBC); Ormstown (JE); Outremont

(UBC). UNITED STATES: Alabama: Baldwin Co.: Bay Minette

(UGA); Colbert Co.: Wilson Dam (LACM); Lee Co.: Auburn

(UWI); Mobile Co.: Mobile (AMNH). Arizona: Coconino

Co.: Flagstaff (OSU); Manzanita Camp, Oak Creek Canyon,

5000' (ASU); Oak Creek Canyon 24 mi S Flagstaff (UAZ);

Walnut Canyon, 6500', 6 1/3 mi ESE Flagstaff (CU);

Williams (OHS); Cochise Co.: Chiricahua Mts, various loaclities (AMNH, ASU, CAS, OHS, UCB, UCR); Douglas (AMNH);

Huachuca Mts (KU, UAZ, YLU); 5 mi W Portal, S. W. R. S.

(AMNH); Sacaton (UCB); 20 mi NE Wilcox (AMNH); Gila Co.:

Sierra Ancha Mts (KU); Greenlee Co.: 6 mi S Gray's Peak

(OSU); Mohave Co.: Hualapai Mt Park (UID); Navajo Co.:

8 - 15 mi NE Whiteriver (AMNH, ASU); Pima Co.: Pepper

Sauce Canyon (CAS); Santa Cruz Co.: Nogales (CAS, UAZ);

Patagonia (CAS, KU, UAZ); Yavapai Co.: Prescott (AMNH,

OHS). Arkansas: Arkansas Co.: vic Stugart (INH); Hot

Springs Co.: Hot Springs (INH); Malvern (MIS); Polk Co.

(KU); Randolph Co.: Imboden (OHS). California: Alameda 91

Co.: Berkeley (CAS, LACM, OSU, UCB); Niles (KU, OSU);

Oakland Hills (CAS, OSU, UCB); Pleasanton (OSU, UCB);

Eldorado Co.: Fallen Leaf Lake (UCB); Humboldt Co.:

Orlock (CAS); Inyo Co.: Bishop (CAS); Lassen Co.:

Susanville (LACM); Los Angeles Co.: Claremont (UMI);

Santa Catalina Island (LACM); 2.5 mi SSW Valyermo, 4800'

(OSU); Plumas Co.: Chester (OHS); Meadow Valley 3500 -

4000' (CAS); San Diego Co.: San Diego (CAS, OSU); San

Francisco Co.: San Francisco (AMNH, CAS); San Joaquin Co,:

Linden (UCR); Vernalis (OSU); San Luis Obispo Co.: Morro

Bay State Park (WSU); Oceana Beach (UCB); Santa Clara Co.:

Cupertino (OSU); Mt View (UAZ); Palo Alto (CAS, CMNH);

Stanford Univ (CAS, LACM); Shasta Co.: Lassen Nati Park

(CAS); Siskiyou Co. (CAS). Colorado: El Paso Co.:

Colorado Springs (CSU); Larimer Co.: Fort Collins (KU);

30 mi N Fort Collins (LSU). Connecticut: Fairfield Co.: Greenwich (AMNH); New Canaan (AMNH); Wilton (AMNH); Kent

Co.: E Greenwich (YLU); Litchfield Co.: Lakeville (UMI);

New Haven Co.: New Haven (CU, YLU); S Meriden (CAS, YLU);

Wallingford (UCT); Woodbridge (YLU); Tolland Co.: Coventry

Twp (UCT); Mansfield (UCT); Stafford Twp (UCT); Storrs

(ASU, UCT). Delaware: Sussex Co.: Seaford (OKS).

District of Columbia:Rock Creek (CU). Florida: Alachua

Co.: Gainesville (CU, UKY); Broward Co.: Hallendale (AMNH); Dade Co.: 3.5 mi W Florida City (UGA); Palm

Beach Co.: Belle Glade (KU); Lake Worth (YLU); Palm 92

Beach (KU); 8 mi NW South Bay (UMO); Sarasota Co.: Sara- sota (PDU); Taylor Co.: Blue Springs Lake (UGA); Georgia:

Bibb Co.: Macon (UGA); Chariton /Clinch Co.: Okefenokee

Swamp (KU); Clarke Co.: Athens (UGA, TAM); UGA Bot Gardens

(UGA); Whitehall Forest (UGA); Clarke Co. (UGA); DeKalb Co.:

Stone Mt (UGA); Hall Co.: Hwy 52, 4.9 mi NW Jct 23 (UGA);

White Sulfur Springs (UMI); Lumpkin Co.: 15 mi N Dahlonega

(UGA); Paulding Co.: Dallas (NDSU); Rabun Co.: Black

Rock Mt State Park (OSU); Seminole Co.: Crk 1.7 mi E Jct

Hwy 39 & 374 (OSU); Tift Co.: Tifton (UGA); Towns Co.:

Hiawassee (UGA). Idaho: Nez Perce Co.: Lapwai (OHS).

Illinois: Adams Co.: Quincy (INH); Champaign Co.: Cham- paign (INH, UMI); Mahomet (INH); Philo (INH); St. Joseph

(INH); Urbana (INH, UCB, UCT); White Heath (INH); Coles

Co,: Charleston (INH); Lincoln Log Cabin State Park (UID);

Cook Co.: Chicago (AMNH, CAS, CU, OSU, UMI, UMO, UNL, WSU);

La Grange (INH); North Chicago (UMI); Edgar Co.: Paris

(INH); Hardin Co.: Elizabethtown (INH); Herod (INH);

Henderson Co.: Oquawka (INH); Iroquois Co.: Watseka (INH);

McHenry Co.: Algonquin (INH); Peoria Co.: Peoria (INH);

St.Clair Co.: E St. Louis (INH); St. Clair Co. (INH);

Sangamon Co.: Carpenter Park (ISM); Springfield (AMNH;

ISU); Union Co.: Anna (INH); Bald Knob (ISM); Cobden (INH);

Vermillion Co.: Danville (INH, MIS); Muncie Ox-bow (INH);

Vermillion Co. (INH); Wabash Co.: Mt Carmel (INH); Winne- bago Co.: Rockford (WSU); Rockton (INH). Indiana: 93

Benton Co. (PDU); Clark Co. (PDU); Jackson Co.: Jackson

Co. Forest Nursery (PDU); Knox Co.: Vincennes (UMI); Knox

Co. (PDU); La Porte Co. (PDU); Monroe Co.: Bloomington

(TAM); Morgan Co. (PDU); Newton Co. (PDU); Perry Co.

(PDU); Porter Co.: Valparaiso (VPI); Posey Co. (PDU);

Steuben Co. (PDU); Tippecanoe Co.: Lafayette (OSU, PDU,

UGA, UMI, UM0); W Lafayette (JE); Tippecanoe (NDSU, PDU

UMI, UMO); Vanderburgh Co.: Evansville (UGA); Vanderburgh

Co. (PDU); Vigo Co. (PDU). Iowa: Audubon Co.: Audubon

(ISU); Buchanan Co.: nr Independence (INH); Clinton Co.:

De Witt (ISU); Dallas Co.: Adel (ISU); Des Moines Co.:

Burlington (ISU, LSU); Dickinson Co.: Cayler Prairie

(TAM); 5 mi W Milford (TAM); Silver Lake Fen (TAM); Jackson

Co.: Maquoketa (LSU, WSU); Lee Co.: Donnelson (ISU);

Fort Madison (ISU, LSU); Linn Co.: Cedar Rapids (ISU);

8 mi E Marion (ISU); Lyon Co.: Little Rock (ISU, UAZ);

Muscatine Co.: Muscatine (ISU); Page Co.: Braddyville

(LSU); Clarinda (ISU); Shenandoah (LSU, MIS, NDSU, TAM);

Pottawattamie Co.: Oakland (ISU); Story Co.: Ames (AMNH, CSU, ISU, KU, LSU, MIS, OHS, OSU, TAM, UCB, UID, UWI, WSU);

Van Buren Co.: Farmington (ISU); Keosauqua (ISU); Woodbury

Co.: Sioux City (ISU, UMN). Kansas: Allen Co.: Iola

(UMN); Bourbon Co. (KU); Clay Co.: Morganville (UCR);

Crawford Co. (UMN); Doniphan Co. (KU); Douglas Co.:

Lawrence (ISU, LACM, OSU); Douglas Co. (KU, LACM, OSU, UMN,

UMO); Gove Co., 2813' (KU); Johnson Co.: Merriam (KU); 94

Johnson Co. (KU); Lyon Co. (KU); McPherson Co. (KU); Miami

Co. (KU); Montgomery Co. (KU); Pottawatomie Co.: Onags

(UAZ); Riley Co.: Manhattan (AMNH, MIS); Shawnee Co.:

Topeka (LACM); Sheridan Co., 2560' (KU); Thomas Co., 3150'

(KU); Woodson Co.: Yates Center (OSU). Kentucky:

Caldwell Co.: Princeton (UKY); Christian Co. (AMNH);

Fayette Co.: Lexington (UKY); Fayette Co. (UKY); Fleming

Co. (UKY); Hart Co.: 2 mi W Horse Cove (UCB); Henderson

Co.: Henderson (UKY); Jackson Co.: Jackson (UKY); Jeffer- son Co.: Louisville (CU); nr Louisville (AMNH); Jessamine

Co. (UKY); Mason Co.: Maysville (UKY); McCracken Co.:

Paducah (UKY); Scott Co.: Georgetown (MIS); Newport (AMNH);

Woodford Co. (UKY). Louisiana: Acadia Par.: Crowley

(LSU); Allen Par. (JE); Baton Rouge Par.: Baton Rouge

(LSU); University (LSU); Caddo Par.: Shreveport (AMNH,

LSU); Cameron Par.: Lacassine (INH); Cameron Par. (INH);

De Soto Par.: Logansport (ISU); E Baton Rouge Par.: Baton Rouge (JE, LSU); Jefferson Par.: Deer Park (LSU);

Harahan (AMNH, WSU); New Orleans (OHS); Shongaloo (LSU);

Madison Par.: Tallulah (UWI); St. Landry Par. (INH, JE);

St. Mary Par.: Franklin (LSU); Tangipahoa Par.: Poncha- toula (LSU); West Feliciana Par. (JE). Maine: Penobscot

Co.: Orono (CAS, MIS, UAZ); not located: Town-ship A

Range 7 (INH); Wales (CAS). Maryland: Baltimore Co.:

Baltimore (AMNH, MSU, VPI); Calvert Co.: Chesapeake Beach

(CU); Port Republic (AMNH); Caroline Co.: Bridgeton (UCT); 95

Frederick Co.: Plummers Lake (CU); Frederick Co. (AMNH);

Montgomery Co.: Cabin John Brown (CU); Prince George's

Co: Beltsville (AMNH); Bladensburg (CU); Bowie Beltwoods

(UCB); Hyattsville (CU); Marsh Hall (CU); Washington Co.:

Hagerstown (CAS); not located:Forest Glen (CU); Maryland

(CMNH, CU). Massachusetts: Barnstable Co.: Provincetown

(AMNH); Woods Hole (AMNH, CAS); Berkshire Co.: Mt Greylock

(CAS); Essex Co.: Pigeon Cove (AMNH); Saugus (CAS);

Swampscott (CAS); Hampden Co.: Chicopee (CU); Hampshire

Co.: Amherst (MIS, MSU, UGA, UMO, VPI); Northampton (CAS,

JE); Ware (AMNH); Middlesex Co.: Framingham (AMNH, CAS, UAZ); Holliston (AMNH, UAZ); Lexington (AMNH); Malden

(AMNH, CU, UKY); Melrose Heights (UCR); Revere (MSU); Sherborn (AMNH, KU); Winchester (UAZ); Nantucket Co: Nan- tucket (INH); Norfolk Co.: Foxboro (VPI); Milton (AMNH,

UMI); Wellesley (AMNH); Suffolk Co.: Boston (CAS);

Worcester Co.: Southboro (AMNH). Michigan: Allegan Co.:

Allegan (MIS); Fennville (MIS); Barry Co.: Hickory Corners

(MIS); Benzie Co. (UMI); Berrien Co.: E. K. Warren Pre- serve (UMI); Gallen (MIS); Herbert Dunes (UMI); New Buffalo

(UMI); St. Joseph (MIS); Berrien Co. (MIS); Cheboygan Co.:

Cheboygan (UMI); Douglas Lake (UMI); Cheboygan Co. (MIS);

Delta Co.: Escanaba (MIS); Eaton Co.: Charlotte (MIS);

Olivet (UGA); Eaton Co. (MIS); Gladwin Co. (MIS, UMI);

Gratiot Co. (UMI); Hillsdale Co.: Cambria (MIS); Pittsford

(MIS); Reading (MIS); Huron Co.: Point aux Barques (UMI); 96

Port Austin (UMI); Sand Point (UMI); Huron Co. (UMI);

Ingham Co.: Aurelius (MIS); E Lansing (MIS, UKY); Mason

(MIS); Millsville (MIS); Stockbridge (UMI); Ingham Co.

(MIS); Tosco Co. (MIS); Isabella Co. (MIS); Jackson Co.:

Francisco (UMI); Jackson (MIS); Kalamazoo Co.: Climax

(MIS); Fulton (MIS); Gull Lake Biol Stn (MIS); Portage

(MIS); Kent Co.: Alto (MIS); Grand Rapids (MIS); Lapeer

Co.: Deerfield Twp (UMI); N Branch (MIS); Lapeer Co.

(MIS); Leelanau Co. (MIS); Manistee Co.: Manistee (UMI);

Manistee Co. (MIS); Monroe Co.: Stony Crk (UMI); Monroe

Co. T8S R6E Sec28 (MIS); Oceana Co.: Shelby (MIS); Silver Lake State Park (UMI); Roscommon Co. (UMI); Schoolcraft

Co. (UMI); Shiawassee Co.: Owosso (MIS); Van Buren Co.: Bangor (MIS); Grand Jot (MIS); S Haven (MIS); Washtenaw

Co.: Ann Arbor (UMI, UMN); Chelsea (MIS); Pittsfield Twp (UMI); Whitmore Lake (AMNH); Washtenaw Co. (UMI).

Minnesota: Brown Co. (UMN); Douglas Co.: Alexandria, lake W of town (UMN); Fillmore Co.: N of Spring Valley (UMN); Freeborn Co. (UMN); Goodhue Co. (UMN); Hennepin

Co.: Minneapolis (UMN); Minnehaha Crk (UMN); Hennepin Co.

(UMN); Lake Co.: Two Harbors (UMN); Mille Lacs Co. (UMN);

Norman Co. (UMN); Ramsey Co.: St. Anthony Park (UMN); St.

Paul (UMN); White Bear (UMN); Rice Co.: Faribault (UMN);

Sibley Co. (UMN); Steele Co.: Medford (UMN); Owatonna

(UMN); Steele Co. (UMN); Yellow Medicine Co. (UMN). 97

Mississiplpi: Grenada Co.: White Lake (LSU); Harrison Co.:

Biloxi (TAM); Handsboro (UMI); Jackson Co.: Ocean Springs

(CAS); Jones Co.: Laurel (TAM); Leake Co.: Carthage

(TAM); Sharkey Co.: INTSOY (INH); Tallahatchie Co.:

Charleston (ISU). Missouri: Atchison Co.: Rockport (UMO);

Tarkio (UMO); Atchison Co. (OSU); Bollinger Co.: Lutesville (UMO); Boone Co.: Ashland (UMO); Columbia (CU, UGA, UMO);

Hartsburg (LSU); Little Dixie (UM0); Rocheport (UM0);

Carroll Co.: Carrollton (UM0); Carter Co.: Big Spring

State Park (MIS); Clinton Co.: Plattsburg (UMO); Gentry

Co.: Albany (UM0); Stanberry (UMO); Hickory Co. (UMO);

Howard Co.: Fayette (UMO); Howard Co. (UMO); Jasper Co.:

Avila (UMO); Livingston Co.: Avalon (UMO); Chillicothe

(UM0); Maries Co.: Vichy (UMO); Morgan Co.: Lake Ozark

(UM0); Ralls Co.: New London (UMO); Perry (UMO); Ste.

Genevieve Co.: 7 mi S Ste. Genevieve (UMO); Vernon Co.:

Nevada (UMO); Vernon Co. (UMO). Nebraska: Antelope Co.:

Neligh (UNL); Blaine Co.: Halsey (UNL); Buffalo Co.:

Gibbon (UNL); Cass Co.: South Bend (UNL); Cuming Co.:

West Point (UNL); Dakota Co.: S Sioux City (UNL); Dawes

Co.: Pine Ridge (UNL); Dixon Co.: Emerson (UNL); Douglas

Co.: Omaha (INH, UNL); Valley (OHS); Lancaster Co.:

Lincoln (OSU, UID, UNL); Roca (UNL); Madison Co.: Norfolk

(UNL); Richardson Co.: Rulo (UNL); Scottsbluff Co.:

Scottsbluff (UNL); Sioux Co.: Sowbelly Canyon (UNL); Sioux Co. (UNL); York Co.: Henderson (UM0). 98

Nevada: Washoe Co.: Verdi (UID). New Hampshire: Grafton Co.: Littleton (AMNH); Plymouth (AMNH); Sullivan Co.:

Claremont (AMNH); not located: Crawfords (CAS). New Jersey: Bergen Co.: Alpine (AMNH); Bear Swamp nr Ramsey

(AMNH); Mahwah (UGA); Palisades (AMNH); Ramsey (AMNH);

Essex Co.: Newark (AMNH); Nutley (AMNH); Gloucester Co.:

Glassboro (INH); Mercer Co.: Fort Dix (OSU); Trenton

(AMNH, UBC); Middlesex Co.: Colonia (ISU); Jamesburg (KU); New Brunswick (AMNH, INH); Old Bridge (AMNH); Morris

Co.: Chester (AMNH); Madison (AMNH); Morristown (JE);

Ocean Co.: Lakehurst (AMNH, CU); Long Beach Island (INH);

Pt Pleasant (CAS, KU); Passaic Co.: Mt View (AMNH);

Passaic Jct (AMNH); Paterson (AMNH); Somerset Co.: Basking Ridge (UCT); Watchung Reservation (UCT); Union Co.:

Elizabeth (AMNH); Rahway (LSU); Roselle Park (UCB). New

Mexico: Dona ASa Co.: Las Cruces (AMNH); Hidalgo Co.:

Red Rock (AMNH); Sandoval Co.:Jemez Springs (USNM); San

Miguel Co.: Las Vegas (AMNH). New York: Cayuga Co.: N

Fair Haven (CU); Springlake (CU); Chautauqua Co.: James- town (UMO); Erie Co.: Buffalo (AMNH, CU, INH, MSU);

Gowanda (CAS); Hamburg (CAS, MSU); Lancaster (ISU, UCR);

Essex Co.: Moriah Center (OSU); Mt Maclntyre (CU); Jefferson Co.: Carlton, Pecton Island (AMNH); Livingston

Co.: Dansville (UM0); Monroe Co.: Honeoye Falls (CU); Mandon (CU); Rochester (AMNH, NDSU, OHS); Rochester Jct

(CU); Niagara Co.: Elma (CAS); Niagara (CU); Niagara Falls 99

(AMNH); Richmond Co.: Staten Island (AMNH, KU); Rockland

Co.: West Nyack (AMNH); St. Lawrence Co.: Clinton (YLU);

Parishville (UMI); Wanakena (AMNH); Suffolk Co.: Calverton

(CU); Cold Spring Harbor (CAS); Coram (AMNH); E Hampton

(KU); E Marion (CU); Fire Island (AMNH); Greenpoint (CU);

Huntington (AMNH); Mattituck (CU); Montauk (AMNH, CU);

Northport (LACM); Orient (AMNH, CU); Riverhead (CU); Smith- town (AMNH); Southampton OSU); Wading River (AMNH); Wyan- danch (AMNH); Yaphank (AMNH, UCT); Tompkins Co.: Beebe

Lake (CU); Ellis Hollow (CU); Freeville (CU); Ithaca (AMNH,

CAS, CU, ISU, KU, OHS, PDU, UMI, UMN); Six Mile Crk (CU);

Verna (CU); Tompkins Co. (CU); Wyoming Co.: Castile (CMNH);

Yates Co.: Himrod (UMO). North Carolina: Bertie Co.:

Lewiston (NCSU); Burke Co.: Morganton (UCB); Carteret Co.:

Beaufort (NCSU); Duplin Co.: Faison (NCSU); Granville Co.:

Oxford (NCSU); Guilford Co.: Greensboro (NCSU); Haywood

Co.: Crestmont (UMI); Macon Co.: Highlands (NCSU); Otto

(UGA); Moore Co.: Eagle Springs (NCSU); Southern Pines (AMNH, CAS, KU); West End (NCSU); Scotland Co. (NCSU);

Wake Co.: McCullers (NCSU); Raleigh (AMNH, CAS, NCSU);

Wake Co. (NCSU); Yadkin Co.: NE of Yadkinsville (NCSU).

North Dakota: Grand Forks Co.: Northwood (NDSU). Ohio:

Ashtabula Co.: Rock Crk (AMNH); Ashtabula Co. (YLU);

Athens Co.: Athens (UCT, UMN, WSU); Athens Co. (YLU);

Butler Co.: Oxford (NCSU); Butler Co. (YLU); Darke Co.

(YLU); Fairfield Co. (YLU); Franklin Co.: Bendon (OSU); 100

Columbus (OHS, OSU, UCT, UMN, WSU); Franklin Co. (YLU);

Fulton Co. (YLU); Hardin Co.: McGuffey (WSU); Hocking Co.

(YLU); Lorain Co.: Amherst (TAM); Lorain Co. (YLU);

Medina Co.: Medina (WSU); Miami Co.: Troy (NDSU); Medina

Co. (YLU); Montgomery Co. (YLU); Ottawa Co.: Lakeside

(UMI); Oak Harbor (WSU); Ottawa Co. (OHS, YLU); Portage

Co.: Kent (CU); Stark Co.: Canton (UMI); Washington Co.:

Marietta (MIS). Oklahoma: Carter Co.: Ardmore (OKS, UNL);

Choctaw Co.: Hugo (OKS); Cleveland Co.: Lexington (OKS);

Norman (OKS); Delaware Co.: Grove (OKS); Delaware Co.

(OKS); Jackson Co.: Blair (OKS); Latimer Co.: Wilburton

(ISU, OKS); Lincoln Co. (OKS); Logan Co. (OKS); Noble Co.:

Marland (OKS); Perry (OKS); Noble Co. (OKS); Nowata Co.:

6 mi N Childers (OKS); Payne Co.: Lake C. Blackwell (OKS);

9 mi W Perkins (OKS); Stillwater (OKS); Payne Co. (OKS);

Pittsburg Co. (UM0); Pushmataha Co.:Antlers (OKS).

Oregon: Benton Co.: Alsea Mt (OSU); Coffin Butte, 10 mi N Corvallis (OSU); Corvallis (OSU); Willamette River (OSU);

Coos Co.: Fairview, 5 mi NE Coquille (OSU); Curry Co.:

Humbug State Park, Brush Crk (OSU); Little Redwood Cmpg,,

T4OS R12W Sec29 (GMS); Pistol River (OSU); Deschutes Co.:

Lava Caves, 15 mi S Bend (OSU); N Sister snowfields 9000;

(OSU); Douglas Co.: 6 mi E Kellogg (AMNH); Jackson Co.:

Table Rock (AMNH); Linn Co.: Albany (AMNH); Marion Co.:

Gervais (OSU); N Howell Prairie (OSU); N Salem (OSU);

Multnomah Co.:Portland(ISU, OSU); Sauvie's Island (OSU); 101

Tillamook Co.: 2 mi SE Pacific City (OSU); Washington Co.:

Forest Grove (ISU). Pennsylvania: Allegheny Co.: Edge- wood (CMNH); 2 mi N Indianola (CMNH); Pittsburgh (CAS,

CMNH); Upper St. Clair Twp (UCB); Allegheny Co. (CMNH);

Beaver Co.: 2 mi NE Industry (CMNH); Bedford Co. (CU);

Berks Co.: Lenhartsville (AMNH); Bradford Co.: Wilawana

(UAZ); Centre Co.: Howard (PSU); State College (PSU);

Cumberland Co.: Mechanicsburg (INH); Dauphin Co.:

Harrisburg (INH); Fayette Co.: Ohio Pyles (CMNH); Lehigh

Co.: Allentown (CU); Mercer Co.: Grove City (UMI); Sandy Lake (AMNH); Philadelphia Co.: Chestnut Hill (KU); Phila- delphia (CU); Washington Co.: Finleyville (CMNH);

Westmoreland Co.: Greensburg (KU); Jeannette (CAS, CMNH,

CSU); Powdermill Nature Reserve (CMNH); Westmoreland Co.

(CMNH). South Carolina: Barnwell Co. (INH); Charleston

Co.: McClellanville (NCSU); Greenville Co.: Greenville

(JE); Pickens Co.: Clemson (MIS). South Dakota: Bennett

Co.: Martin (ISU); Brookings Co.: Brookings (CAS, UNL);

Yankton Co.: Yankton (CAS). Tennessee: Blount/Sevier

Co.: Great Smoky Mt5Natl Park (OHS); Carter Co.: Roan Mt,

6106' (UMI); Cumberland Co.: Grassy Cove (UMI); Henderson

Co.: Natchez Trace State Park (MIS); Knox Co.: Knoxville

(ISU); Roane Co.: 10 mi N Oak Ridge (UNL); Sevier Co.:

Administration Area, Great Smoky Mts Natl Park (AMNH).

Texas: Bell Co. (TAM); Brazos Co.: Bryan (TAM); College

Stn (JE, MIS, PDU, TAM) ; Koppe Bridge (TAM) ;Brazos Co. 102

(AMNH, CSU, MIS, TAM); Colorado Co. (KU); Erath Co.:

Stephenville (JE); Galveston Co.: Galveston (WSU); Hardin

Co.: 9 mi W Beaumont (OSU); Kaufman Co.: 2 mi W Forney

(TTU); Lubbock Co.: Airport, Lubbock (TTU); 3 mi W Lub- bock (TTU); McLennan Co.: Waco (AMNH, TAM); Sabine Co.:

Sabine Nati Forest, Lakeview Cmpg (TAM); Travis Co.:

Austin (CAS, CU, PDU); Uvalde Co. (OHS); Williamson Co.:

Taylor (TAM). Utah: Washington Co.: Zion Nati Park (ISU).

Vermont: Franklin/ Lamoille Co.: Green nits (CAS, CSU);

Lamoille Co.: Stowe (AMNH); Orleans Co.: E Charleston

(AMNH); Windham Co.: Putney (AMNH); Windsor Co.: Wood- stock (AMNH). Virginia: Albemarle Co.: Crozet (VPI);

Charlotte Co. (VPI); Chesapeake (VPI); Falls Church (AMNH);

Fauquier Co.: Paris (CU); Giles Co.: Mt Lake Biol Stn,

3900' (KU); Grayson Co. (VPI); King & Queen Co. (VPI);

Montgomery Co.: Blacksburg (VPI); Montgomery Co. (VPI);

Princess Ann Co.: Backbay (UCR); Richmond (VPI); Rock- bridge Co.: Greenville (VPI); Steele's Tavern (VPI);

Rockbridge Co. (VPI); Shenandoah Co.: Woodstock (CU);

Staunton (VPI); Virginia Beach (VPI). Washington: Cowlitz

Co.: Longview (OSU); Franklin Co.: Pasco (UCB); Pacific

Co.: Bay Center (OSU); Seaview (ASU); Pierce Co.:

Paradise Park, 6000' (CAS); Puyallup (WSU). West Virginia:

Barbour Co.: Philippi (CMNH); Harrison Co.: Salem (UMI);

Lewis Co.: Jackson's Mill (CMNH); Logan Co.: 4 mi SE

Logan (OKS); Morgan Co.: Berkeley (CU); Berkeley Springs 103

(CU); Pocahontas/Randolph Co.: Cheat Tits (CMNH); Ritchie

Co.: Harrisville (PDU). Wisconsin: Crawford Co.: Gays

Mills (UWI); Dane Co.: Madison (ISU, UWI, WSU); Dane Co.

(UWI); Door Co. (UWI); Fond du Lac Co. (UWI); Juneau Co.

(WSU); Milwaukee Co.: Milwaukee (KU); Wauwatosa (UWI);

Polk Co.: Amery (UWI); Gibson Lake (UWI); Racine Co.:

Racine (ISU); Rush Co.: Hawkins (UWI); Sauk Co.: Sauk

City (UGA); Sauk Marsh (UGA); Trempealeau Co. (UWI);

Waupaca Co.: Clintonville (WSU); Waushara Co., T2ON R11E

(UWI).

This species was long known as P. spinosus (Dallas),

as Say's (1831) description of Pentatoma maculiventris,

published at New Harmony, Indiana, was not discovered until

1899 and reprinted by Scudder. Dallas' type of Arma

spinosa 1851 is a specimen with spinose humeri which fits

Say's description. Dallas (1851) described Arma modesta

based on a specimen with subacute humeri. The male

genitalia and ventral spine are identical to Arma spinosa

and Podisus maculiventris. The syntype of Arma pallens

St&l that I examined, a female, has a long ventral spine

typical of maculiventris. The humeri are subacute and not

much produced.

This species is most often confused with P. serieven-

tris and P. brevispinus due to the variation in humeral

shape. The latter 2 species never have spinose humeri.

A single series of Podisus maculiventris may vary in the 104

extent to which the humeri are produced. Generally,

smaller individuals have subacute humeri whereas the latter

tend to be acute to spinose in larger specimens.This may be correlated with the number of prey consumed by nymphs

but remains to be tested. Evans (1982b) noted that

later-maturing individuals were smaller due to scarcity of prey, but did not mention humeral variation.

Podisus maculiventris has a longer ventral spine than either P. serieventris or P. brevispinus and is usually

larger. Male genitalia of maculiventris and serieventris, are indistinguishable, but I have not synonymized the 2 species because of differences in humeri, coloration, and ventral spine length. Hybridization is discussed under P. serieventris, p. 120.

Variation exists in the claspers and spermathecal bulb. The lateral edge of the former may be a smooth curve between the dorsal and lateral apices, or form a

100 - 120°angle midway between these points. These differences are random, or not associated with a particular geographical area. The spermathecal bulb is elongate but varies in the amount of curvature. In some individuals the bulb is slightly constricted near the tip.Some of this variation may be due to specimen preparation; however, there is no correlation between bulb shape and particular geographic localities. 105

P. maculiventris, a widespread generalist predator,

has been studied extensively because of its potentialas a

biological control agent. Results of several life history

studies are shown in Table 1. I reared a generation of

this species from eggs sent tome by Dr. K. V. Yeargan,

University of Kentucky, Lexington. The results did not

differ considerably from those of other workers (Table 1).

McPherson (1982) gives a list of 103 prey species

distributed among 43 families in 8 orders.Additions to

this are presented in Table 2. Noctuid larvae are P. maculiventris' most common prey. Among the insects consumed

are serious pests: Lyqus lineolaris (Palisot de Beauvois)

(Miridae), Nezara viridula (L.) (Pentatomidae), Epilachna varivestis Mulsant (Coccinellidae), Leptinotarsa

decemlineata (Say) (Chrysomelidae), Hypera postica

(Gyllenhal) (Curculionidae), Anticarsia qemmatalis HUbner

(), Trichoplusia ni (HUbner) (Noctuidae);

Heliothis virescens (F.) (Noctuidae), H. zea (Boddie)

(Noctuidae), Pieris rapae (L.) (Pieridae), and Diprion similis (Hartig) () (McPherson 1982, references therein).

P. maculiventris was introduced into Europe (Couturier

1938) in an unsuccessful attempt to control the Colorado potato beetle. In Yugoslavia, P. maculiventris and P. placidus, introduced to suppress Hyphantria cunea (Drury), the fall webworm, did not overwinter successfully 106

(Tadi 1975). P. maculiventris was released in the U. S.

S. R. in 1974 for control of Leptinotarsa decemlineata

(Sazonova et al. 1976). These workers concluded that acclimatization should not be difficult.

Japanese efforts to establish this predator for,-sup- pression of H. cunea since 1970 have not been successful

(Hokyo and Kawauchi 1975). Conversely, Kim et al. (1968) reported promising results with the same predator and prey in Korea. Perhaps the unsuccessful predators were imported from areas with environmental conditions unlike those where release occurred. Because P. maculiventris is found throughout a wide range of environments in America north of Mexico, suitable biotypes could probably be found for re-introduction.

Research into various aspects of predation in P. maculiventris is fairly extensive and ongoing. Baker

(1927) described the mouthparts and the manner in which host tissue is penetrated. Cutting is done by serrately tipped mandibles in a sliding motion. The bug is able to maintain its hold on the prey due to inward curling of the stylet tips. When the latter are pressed together they straighten and allow for quick removal from host tissue.

Gallopin and Fitching (1972) estimateda mean in- gestion rate of 1.801 mg / hr / bug which apparently was not affected by initial prey weight, age, or species of 107

prey. This corroborates results of Mukerji and LeRoux

(1969a). However, as initial prey weight increased, so did

ingestion rate. The authors suggested that internal

pressure of prey tissues affects the predator's rate of

suction. Gallopin and Kitching (1972) also found thatas

length of starvation period increased, percentage of

insects feeding after re-exposure to prey declined.

The functional response of P. maculiventris has been

studied by Morris (1963) on Hyphantria cunea (Drury),

Mukerji and LeRoux (1969a) on (L.), and

Waddill and Shepard (1975a) on Epilachna varivestis Mulsant.

All found that as prey density increased, the number of

successful attacks rose at a decelerating rate. Morris

(1963) noted that larval defense affected P. maculiventris'

functional response. The predator consumed 17.4 defense-

less H. cunea larvae in 20 days whereas 10.8 active larvae were eaten over an equal period of time. Predators

ingested about 84% of their own body weight with each prey

consumed and were therefore also limited in functional

response by handling time (consumption and digestion).

The latter, along with attacking time averaged 1.15 days

(Morris 1963). As the predators aged, predation rates decreased (Morris 1963).

Morris (1963) performed his studies considering density of webworms per nest.but felt that in the fields density of nests was important. Also, in a field situation 108 some larvae in each nest would be parasitized, diseased or otherwise weakened, and therefore more easily subdued by

P. maculiventris (Morris 1963).

Morris (1972) observed P. maculiventris, and less commonly, P. brevispinus, P. serieventris, P. placidus, and A. bracteatus, to be important fall webworm predators.

Eggs were found inside fall webworm nests, but the number of predators per nest was small. As Podisus spp. are timid, and averaged 0.54 prey killed per day in his study,

Morris (1972) concluded that these predators were a small but steady mortality factor, especially for medium -sized larvae.

Mukerji and LeRoux (1969b, c) studied food consumption and energetics of P. maculiventris. Nymphs and adults supplied with different amounts of prey varied in rate of capture, consumption, and growth. Nymphal food consumption was linearly related to growth and development. Female adults differed in food intake at each diet level and number of eggs laid. The results (Mukerji and LeRoux

1969b) showed that although the insects could survive for long periods on small prey, feeding on larger prey increased developmental rates and maintained higher fecundity. Studies on energetics showed that growth efficiency was highest in nymphs and lowest in male adults

(Mukerji and LeRoux 1969c). Females used most of their energy for egg production. 109

The phenology of P. maculiventris has been examined relative to 2 prey species, Hyphantria cunea (Morris 1963,

1972) and Trirhabda sp. (Evans 1982a). The former stated that this predator is well-synchronized with H. cunea, while the latter, more extensive work, revealed a lack of synchrony with Trirhabda sp. in Solidago sp. stands. Timing of oviposition by Podisus was seen as a crucial factor in its asynchrony with the chrysomelid species.

Evans (1982a) hypothesized that frequency of encounters between female predators and larval Trirhabda may stimulate oviposition in the former. By the time second instar

Podisus are present, Trirhabda larvae are maturing and beginning to enter the soil for pupation. Podisus nymphs are able to prey on other larvae but also have to spend more time for each successful attack on active Trirhabda adults which begin appearing while the pentatomids are still maturing.

Evans (1982b) found later-maturing Podisus to be smaller than laboratory reared individuals supplied with an abundance of Spodoptera eridania (Cramer). Smaller females had 66% of the fecundity of larger bugs (Evans

1982b). While P. maculiventris is polyphagous, Evans

(1982a) points out that it may actually specialize in seeking outbreaks by ovipositing in a variety of habitats. Although it may stabilize outbreaks, this species rarely prevents them (Evans 1982a). 110

The suitability and effectiveness of P. maculiventris on certain pest species has recently been examined.

Waddill and Shepard (1975a) found that the predator took comparable numbers of Epilachna varivestis Mulsant

(Mexican bean beetle) and Anticarsia qemmatalis Hubner

(Velvetbean caterpillar) as well as a wide variety of stages of the former. The authors concluded that P. maculiventris was opportunistic, as it found suitable prey throughout the season in South Carolina .

In a comparison of carnea Stephens,

Coleomeqilla maculata (DeGeer), Geocoris punctipes (Say), and Podisus maculiventris as predators of Heliothis zea

(Boddie) (corn earworm) and H. virescens (F.) (tobacco budworm), Lopez et al. (1976) found P. maculiventris to be most effective on third instar prey. The authors point out that as few naturally occurring predators attack large

Heliothis larvae, P. maculiventris could be useful in a conservation or augmentation biological control program.

Marston et al. (1978) exposed 5 species of soybean caterpillars to predation by P. maculiventris.Anticarsia, qemmatalis Hubner and Plathypena scabra (F.) exhibited the most vigorous defensive reactions and therefore were least successfully attacked. Those offering minimal resistance,

Trichoplusia ni (Hubner) and Pseudoplusia includens

(Walker), suffered 75 - 80% mortality, while Heliothis zea was intermediate (60% mortality). Richman and Hemenway 111

(1980) evaluated 16 species as predators of P. includens in a field cage study and found that P. maculiventris specialized on medium-sized larvae. Information of this nature is useful in determining which predator is best- suited for controlling injurious insects in a given situation.

One problem with Podisus and other winged predators is dispersal. Ignoffo et al. (1977) found that de-winged female P. maculiventris laid 5 times more eggs on potted soybeans than winged females. In field tests, 85% of de-winged individuals were still present in soybean fields

72 h after release as opposed to 12% of winged bugs.

While this method is not practical for mass release programs, the authors suggest that a genetic means of producing wingless (flightless) P. maculiventris be investigated.

Waddill and Shepard (1975b) point out that dispersal patterns of agents considered for mass release should be ascertained. Their results with P. maculiventris nymphs in soybeans indicate that dispersal was along rather than across rows, although no canopy overlap was present.

An innovative approach to biological control was taken by Biever et al. (1982) who surface-contaminated P. maculiventris adults with Trichoplusia ni nuclear polyhedrosis virus and allowed the insects to walk on filter paper disks or soybean leaves. Larval T. ni 112 mortality rate after exposure to 1- 4 day old contami- nated filter papers was over 90%, whereas 11 day old disks produced a 50% death rate. In field studies, contaminated predators started epizootics which effectively suppressed the pests within 3 - 4 weeks.

For Podisus spp. to be part of an integrated control program, it and any insecticides used would have to be compatible. Wilkinson et al. (1979) tested the effects of 4 synthetic pyrethroids and organophosphate insecticides on several beneficial insects. P. maculiventris nymphs and adults showed 73.8 - 98% survival after exposure to the synthetic pyrethroids fenvalerate and permethrin, but 93-

100% mortality due to the organophosithates sulprofos and profenofos.

Chemical studies of Podisus spp. glandular secretions are in progress. Aldrich et al. (1978) found sexual dimorphism in adult P. maculiventris III -IV dorsal abdominal glands. Males have much larger glands, which the authors felt might be used in aggregation for reproduction or to aid males in overcoming female predatory instincts. The secretion contains a diverse array of chemicals, predominantly (E)-2-hexenal, o(-terpineol, and benzyl alcohol, which implies that aliphatic, terpenoid, and aromatic pathways are involved in scent manufacture.

These compounds are not believed to be derived from food as they are present in large quantities; however, the 113 possibility of symbiont involvement in their manufacture has not been investigated.Aldrich recently has

synthesized sex pheromones for P. maculiventris and P. placidus.

In other chemical studies, McLain (1979) observed that P. maculiventris would follow artificial terrestrial trails made by mixing a small amount of water with frays of fourth instar Malacosoma americanum (F.). The bugs swept or tapped their antennae on the trail and extended their rostra, pausing at intervals to groom these appendages. McLain (1979) speculated that using kairomones to locate prey might be advantageous for predators such as these that have small eyes.

P. maculiventris is attacked by several egg and adult parasitoids (Table 3). Buschman and Whitcomb (1980) observed phoretic behavior in an undescribed , a phenomenon not recorded for heteropteran parasitoids outside of Africa and Sumatra. Parasitoid adults clung most frequently to the female bug's pronotum; occurrence on males was short - lived. Soon after P. maculiventris oviposited, the Telenomus would drop off its host to parasitize the freshly laid eggs. Host specificity was apparent, as the would not mount gravid P. fretus or

Euthyrhynchus floridanus (L.) females (Buschman and

Whitcomb 1980). The possibility that these parasitoids are cuing on a P. maculiventris pheromone remains to be 114 investigated.

Podisus serieventris Uhler

Fig. 9; Map 9)

Podisus serieventris Uhler 1871: 94; Van Duzee 1904: 71.

Apateticus (Eupodisus) serieventris Schouteden 1907: 72.

Apateticus (Podisus) serieventris Kirkaldy 1909: 20.

Podisus (Podisus) serieventris Van Duzee 1917: 79.

TYPE AND TYPE LOCALITY: Podisus serieventris Uhler, type, male, poor condition (MCZ): "40.S / 13 / M. C. Z. Type

26429 (red) / Podisus serieventris; 13. Uhler." Uhler gives Cambridge, Massachusetts as the type locality.

DIAGNOSIS: Coloration gray to brown, mottled with fuscous; ventral spine not extending between hind coxae. Clasper triangular with concave lateral edge.

DESCRIPTION: Body length: male 8.5 - 11.5; female 10.0 -

12.0. Humeral width: male 4.4 - 6.0; female 4.7 - 6.7.

Dorsum yellowish gray to brown, mottled with dark gray to fuscous; punctation brown to fuscous or black; venter yellow with dense fuscous or red punctation. Head densely punctured with fuscous or black; width:male 2.0 - 2.4; female 2.1 - 2.5; length: male 1.7 - 2.2; female 2.1 -

2.3. Antennae brown to fuscous with paler incisures; segments III - V darker than I and II.Pronotum with margins anterior to humeri straight, slightly explanate, 115 ivory, dentate; punctation most dense on humeri and interior to margins. Humeri usually bluntly rounded, not much produced; in some specimens tending toward subacute.

Scutellum: width: male 2.2 - 3.2; female 2.6 - 3.4; length: male 2.9 - 4.1; female 3.4 - 4.5. Membrane yellow with dark apical band. Legs yellow to orange-red with variably defined fuscous dot near femoral apices; punctation ranging from barely discernable to dense, dark; tibiae sulcate dorsally. Ventral spine not or barely reaching hind coxae. Male genitalia: width of genital cup 1.31 - 1.64; very similar to preceding species.

Claspers triangular with concave lateral margins variable as in preceding species. Genital plates quadrate, produced on ventral half of mesal margin; smoothly indented laterally; surface irregularly ridged, tuberculate.

Female genitalia: width of first gonocoxa 0.58 - 0.90; length 0.53 - 0.69; margined posteriorly with fuscous.

Bulb as in P. maculiventris. Length of scierotized rod

0.76 - 1.16.

SPECIMENS ILLUSTRATED: Humerus, genital cup, ventral spine: Michigan: Oceana Co.: Mears (MIS); clasper:

Montana: Gallatin Co. (MSU); spermatheca: Michigan:

Chippewa Co.: 14 mi S Sault Ste. Marie (MIS).

DISTRIBUTION: Across Canada and the United States except in the states east of New Mexico and south of North 116 Dakota, Louisiana, and Georgia (Map 9).

MATERIAL EXAMINED: 349 specimens from the following localities: CANADA: British Columbia: Forbidden Plateau (UBC); Golden Ears (UBC); L. Cowichan (LSU, UCT); Langley

(UBC); Mt Todd (UBC); Pemberton Meadows (UBC); Riske Creek

(UBC); Saanich (UBC); Seymour Mt, 4000' (UBC); Vancouver

(UAZ); Vernon (UBC); Victoria (INH, UBC). Nova Scotia:

Baddeck, Cape Breton (AMNH); Grand River (CAS, ISU, UCT).

Ontario: Brule Lake (CAS); Footes Bay (UCB); 15 mi SE

Kenora (UCB); Lake Temagami (CAS). Quebec: Cascapedia R. (UCT); Ft Coulonge (CAS); Montreal (JE, UBC); Outremont

(UBC). Not located: Beaverfoot Range, Rocky Mts (AMNH).

UNITED STATES: Arizona: Cochise Co: Huachuca Mts (KU);

Rustler Park, Chiricahua Mts 8200' (ASU); Graham Co.:

Pinaleno Mts 8950' (CAS); Pima Co.: Catalina Mts, Ariz 22

HkHy (UAZ); not located: Phelps Bot Area, White Mts 9500'

(CU). California: Eldorado Co.: Fallen Leaf Lake (UCB);

Modoc Co.: Lake City, Warner Mts (CAS); Plumas Co.:

Meadow Valley 3500 - 4000' (CAS). Colorado: El Paso Co.:

Rock Creek Canyon 8500' (AMNH); Ouray Co. (CU); Teller Co.:

Pike's Peak (UCB), Connecticut: Tolland Co.: Willington

Twp, Carly Tract nr Mansfield (UCT); Windham Co.: Putnam

(AMNH); not located: Woodstock (CAS). District of

Columbia: (Cu)*. Georgia: Clarke Co.: Athens Memorial

Park (UGA)*, Idaho: Boundary Co.: Bonners Ferry (UID);

Latah Co.: 1 mi E Deary (UID); Moscow (ISU, UID); Moscow 117

Mt (ISU, UID); Lewis Co.: Lewiston (UID). Illinois:

McHenry Co: Algonquin (INH); not located: Beach, lake shore (INH); "N. ILL." (INH). Indiana: Crawford Co.:

Wyandotte (PDU)*; Fulton Co. (PDU). Kentucky: Christian

Co. (UKY)*. Louisiana: Winn Parish: Winnfield (UMI). Maine: Cumberland Co. (AMNH); Lincoln Co. (OHS); Penobscot

Co.: Orono (CAS); Piscataquis Co.: Baxter State Park

(CU); Mt Katandin (AMNH, UCB); not located: "Maine"

(CAS, CU). Maryland: Prince George's Co.: Bladensburg

(Cu)*. Massachusetts: not located: Beach Bluff (CAS). Michiaan: Alcona Co. (MIS); Charlevoix Co.: Bay Shore

(INH); Cheboygan Co.: Cheboygan (UMI, OSU); Chippewa Co.:

14 mi S Sault Ste. Marie (MIS); Whitefish Pt (UMI);

Crawford Co. (MIS); Emmet Co.: Waucoskancea (UMI);

Houghton Co.: Lake Superior Beach (UMI); Huron Co. (UMI);

Ingham Co.: East Lansing (MIS); Iosco Co.: Oscoda (UMI);

Kalamazoo Co.: Gull Lake Dial Stn (MIS); Keweenaw Co.:

Lake Superior Beach (AMNH); Lake Co. (MIS); Livingston Co.

(UMI); Mackinac Co.: (UMI); Naubinway (UMI);

"Str Search" (UMI); Marquette Co.: Marquette (CU, UMN);

Oakland Co.: Milford (UMI); Oakland Co. (UMI); Oceana Co.:

Mears (MIS); Schoolcraft Co.: Thompson (KU); Schoolcraft

Co. (MIS); Washtenaw Co.: Ann Arbor 3rd Woods (UMI);

Washtenaw Co. (UMI). Minnesota: Cook Co.: Granite Lake 50 mi NW Grand Marais (UNL); Crow Wing Co.: Mille Lacs

(UMN); Goodhue Co. (UMN); Olmstead Co. (UMN); St. Louis 118

Co.: Duluth (INH, UMN); not located: Eaglesnest (INH);

"Min" (INH); Shevlin (KU). Missouri: Boone Co.: Ashland

Wildlife Area (UMO) *; Callaway Co.: Holts Summit, Camp

Keown (DBT)*; St. Louis Co.: Ranken (UMO) *. Montana:

Fergus Co.: 6 mi N Lewistown (MSU); Gallatin Co.:

Bozeman (MSU); Gallatin Co. (MSU); Jefferson Co.: Nigger

Hollow (MSU); Lake Co.: Swan Lake (UID); Sanders Co,:

White Pine (MSU); not located: Lake Roan (LSU). New

Hampshire: Coos Co.: Mt Washington (AMNH, CAS, INH,

YLU); Grafton Co.: Franconia (AMNH); Sullivan Co,:

Claremont (KU); not located: Three Mile Island (AMNH);

White Mts (AMNH). New Mexico: Catron Co.: Mogolion Mts

(UCB); Santa Fe Co.: Tesuque (AMNH, ASU); not located

White Mts, S Fork Eagle Creek 8000' (CSU). New York:

Clinton Co.: Peru (CU); Cortland Co.: Heiberg Forest

(UMI); Erie Co.: Clarence (ISU); Essex Co.: Heart Lake

(CU); Lake Placid (CAS); Mt Maclntyre (CU); Mt Whiteface

(CU); New Russia (CU); Franklin Co.:Malone (CU); Greene

Co.: Maplecrest, Catskill Mts (AMNH); Greene Co. (CMNH);

Livingston Co.: Hemlock Lake (CU); Orange Co.: Bear Mt

(AMNH); Huguenot (AMNH); Schuyler Co.: Texas Hollow (CU);

Suffolk Co.: Bayshore (AMNH); Montauk (CU); Orient (CU);

Tompkins Co.: Ithaca (AMNH, CU). North Carolina:

Buncombe Co.: Black Mt, N Fork Swanannoa (AMNH); Watauga

Co.: nr Boone (NCSU). North Dakota: Cass Co.: Fargo

(NDSU). Ohio: Greene Co. (OHS); Hocking Co. (OHS, YLU). 119

Oregon: Benton Co.: Corvallis (CAS); Kings Valley (LSU);

Douglas Co,: Diamond Lake (AMNH); Jackson Co.: Union

Creek (OSU); Klamath Co.: Chiloquin, Chiloquin Ridge

(OSU); Lane Co.: H J Andrews Exp Forest (DBT); McKenzie

Bridge (OSU); Triangle Lake (LSU); Linn Co.: nr Cascadia, Trout Creek Forest Camp (OSU); House Rock Cmpg (OSU);

Sweet Home (OSU); Union Co.: La Grande (OSU).

Pennsylvania: Bedford Co.: New Paris (CMNH); Bedford Co.

(CU); Centre Co.: Benner Springs (PSU); Livonia (PSU);

State College (PSU); Erie Co,: Lake Erie, S Shore (AMNH);

Huntingdon Co.: Neff Mills (PSU); not located: Colerain

Park (PSU); Galbraith Gap (PSU); Gap Run (PSU). Utah:

Garfield/Kane Co.: Bryce Canyon (CAS); Utah Co.: Aspen

Grove Camp, Mt Timpanogos (ISU). Vermont: Lamoille Co.:

Stowe (AMNH); Orleans Co.: E Charleston (CU); Windsor

Co.: White River Jct (JE). Virginia: Falls Church

(AMNH) *; Herndon (UMI) *. Washington: Asotin Co.: Fields Spring State Park, 4 mi S Anatone 3500 - 4000' (WSU);

King Co.: North Bend (OSU); Roy (UCB); Seattle (CAS);

Klickitat Co.: Bowman Creek, 12 mi NW Goldendale (TAM);

Snohomish Co.: Monroe (CAS); Stevens Co.: Deer Lake nr

Chewelah (WSU); Thurston Co.: Olympia (OSU); Whatcom Co.:

Mt Baker (KU); not located: Cliffdell (KU); Maywood

(OSU). Wisconsin: Dodge Co.: Beaver Dam (CAS); Polk

Co.: Gibson Lake T34N R16W Sec34 (UWI); Vilas Co.:

Northern Highland (UWI). Wyoming: Albany Co.: Curtis 120

Gulch Cmpg T28N R73W Sec8 (INH); Medicine Bow Nati Forest

(MIS).

Prebble (1933) referred to color variations in P. serieventris, an observation I have also made. Adults range from grayish brown to blackish gray with reddish to pale anterolateral margins.

This species is often confused with P. maculiventris and P. brevispinus. The latter differs from serieventris in the shape of its claspers and humeri (fig. lla, c) as well as its lighter coloration. Differences between P. serieventris and P. maculiventris are discussed on pages

103 - 104.

Hybridization between serieventris and maculiventris occurs in the laboratory. Coppel and Jones (1962) observed copulation between a female P. maculiventris and a male P. serieventris which resulted in viable progeny. The authors reared these offspring through 4 generations and described them as intermediate in humeral width and egg characteristics. Total nymphal development time was closer to that ofmaculiventris. Voucher specimens could not be located (H. C. Coppel, pers. comm.).

I have examined specimens that are difficult to place in either species, most often because of a longer ventral spine than is common in serieventris combined with the typical coloration of maculiventris. These individuals usually have humeri intermediate between the 2 species, as 121 described by Coppel and Jones (1962).Specimens examined and found to have intermediate characteristics are marked with an asterisk (*).

Most intermediate individuals were from the southern portions of serieventris' range.This area roughly corresponds to Remington's (1968) Southern Appalachian -

Ozark - Ouachita Suture Zone. The presence of this zone is attributed to a break in the forest which once covered the area, allowing species typical of open areas to interbreed with related forest-dwelling taxa.The extent of hybridization and documentation of its occurrence naturally in these two species needs to be investigated.

Kirkland (1896) described the eggs, nymphs, adults, and the life history of this species, but recorded only 4 instars. All subsequent studies on this and other Podisus species have reported 5 instars. Prebble (1933) studied the biology of P. serieventris, described immature stages, and reported the insect's role in suppressing an outbreak of Ascieris variana (Fernald), the Eastern blackheaded budworm, in Nova Scotia. McPherson (1982) lists 39 insect species that are preyed upon by P. serieventris. I have recorded additions to the latter's list in Table 2. Reeks (1938) reported that P. serieventris was the most common predator attacking larvae of Gilpinia hercyoniae (Hartig), the European spruce sawfly, in

Canada. Forty-eight predators and 1009 sawfly larvae were 122 beaten from 1 tree in Fredericton, New Brunswick. Nymphs were collected between July 20 and August 31, whereas adults were taken from August 24 to September 21.

Coppel and Jones (1962) studied the species' bionomics in Wisconsin. Results of these studies are shown in Table 1. Trissolcus podisi Ashmead (Hymenoptera: Scelionidae) parasitizes the eggs of P. serieventris (Table 3)

(Krombein et al. 1979).

Podisus conarex (Stal)

(Fig. 10; Map 10)

Telepta conarex StAl 1862: 91.

Arma conclrex Walker 1867: 135.

Podisus (Podisus) conarex Stal 1870: 51.

Podisus conarex Distant 1880: 40, pl. 1, fig. 20.

Apateticus (Eupodisus) conarex Schouteden 1907: 71.

Apateticus (Podisus) conarex Kirkaldy 1909: 18.

TYPE AND TYPE LOCALITY: Telepta, conarex Stal, 2 syntypes, both in excellent condition (Riksmuseet): female:

"Mexico / Sal14. / Type. / Typus (red) / 410; 82 (red) /

Riksmuseum; Stockholm (green);" male: "Mexico / Salle. /

Type / Allotypus (red) / 411; 82 (red) / Riksmuseum;

Stockholm (green)." The red type labels are for museum purposes only and are of no nomenclatural significance 123

(P. Lindskog, pers. comm.). The museum invoice lists the specimens as syntypes. I will propose that the male be given lectotype status in a subsequent publication. Seal gives no type locality in his original description, but names Mexico in the 1870 reference.

DIAGNOSIS: Anterolateral margins of pronotum swollen, polished, dentate; humeri acute, produced laterally, tipped with red or fuscous. Claspers as in fig. 10c, d.

DESCRIPTION: Body length: male 10.0 - 11.0; female 10.5 -

13.5. Humeral width: male 5.8 - 7.4; female 6.4 - 8.2. Dorsum light brown with yellow wash, brown punctation; venter pale yellow, polished; punctation concolorous.

Head fuscous punctate; width: male 2.0 - 2.3; female 2.2 -

2.5; length: male 1.7 - 1.9; female 1.8 - 2.2. Antennae ochraceous; segments III, IV, V with apical half reddish brown, basal half pale. Pronotum with dentate anterolateral margins tumid, ivory, polished. Humeri densely punctate, produced, acute; tips polished red t fuscous. Scutellum: width: male 2.5 - 3.0; female 2.9 3.6; length: male 3.3 - 3.8; female 3.7 - 4.8; median pale line extending from base to apex. Membrane pale golden with dark apical band. Legs concolorous with venter; tibiae not sulcate dorsally; tarsi darker. Venter sometimes with lateral row of small fuscous dots. Ventral spine extending between hind coxae. Male genitalia: 124 width of genital cup 1.12- 1.38; lateral lobes broadly rounded, entire margin setose. Claspers triangular; dorsal portion narrow, small; lateral concavity short, shallow, allowing for deep, blunt basal portion with toothed dorsolateral tip. Genital plates quadrate, dorsally and medially convex; surface with flattened tubercles. Female genitalia: Width of first gonocoxa

0.48 - 0.66; length 0.42- 0.53; Bulb ovate, flanges prominent. Length of sclerotized rod 1.09 - 1.19.

SPECIMENS ILLUSTRATED: Humerus, spermatheca: MEXICO:

San Luis Potosi: 30 mi S Tamazunchale (CAS); genital cup:

ECUADOR: Banos (UCB); clasper: ECUADOR: Banos (UCB).

DISTRIBUTION: Mexico (San Luis Potosi and southward),

Guatemala, Costa Rica, Haiti, Colombia, Ecuador.

MATERIAL EXAMINED: 47 specimens from the following localities: COLOMBIA: Magdalena: Socorpa Mission, Sierra de Perija (AMNH); Pueblo Bello, 45 km W Valledupar,

Sierra Nevada de Santa Marta (AMNH); Espinal (AMNH).

COSTA RICA: CArtago: Pacayas (AMNH). Puntarenas: nr

San Vito (VPI); Santa Elena de Monteverde (DBT). ECUADOR:

Banos (UCB); El Tabldn (UCB); Mt Tunguarahua (UCB); Rib

Blanco (UCB); Runtlin (UCB). GUATEMALA: Guatemala (AMNH);

Nebaj (AMNH). HAITI: La Visite, La Selle Range 5 - 7000'

(AMNH). MEXICO: Jalisco: Nevado de Colima Rd 5.9 mi W

Hwy Jct nr Atenquique (JE). Mexico: 7 mi N Acambay (TAM). 125

Morelos: Cuernavaca (CAS, PSU). Oaxaca: 13 mi S

Suchixtepec (TAM). San Luis Potosi: 30 mi S Tamazunchale

(CAS). Veracruz: Fortin de las Flores (UCB); Rio Blanco

(UCR).

Podisus congrex is a Neotropical species found from San Luis Potosi, Mexico south through Central America and into Colombia and Ecuador. Individuals may stray northward into the Nearctic zone and for this reason I have included it in this treatment.

The biology of P. congrex is unstudied.

Podisus brevispinus, new species

(Fig. 11; Map 11)

Podisus modestus, authors not Dallas 1851.

TYPE AND TYPE LOCALITY: Holotype, male, excellent condition, to be deposited in the U. S. National Museum:

"Dane Co., Wis; May 30, 1911; Wm. S. Marshall / Collection of Wm. S. Marshall." Paratypes (54): CANADA: British

Columbia (14): Saanich District, various dates, 6 males,

8 females (UBC). UNITED STATES: Colorado (8): Larimer

Co.: Fort Collins, various dates (CSU, KU, UAZ). New

Hampshire (5): Coos Co.: Bretton Woods, June, 30, 1909,

3 males, 2 females (CAS); New Jersey (6): Berkeley

Heights, various dates, 3 males, 3 females (AMNH). North

Carolina (6): Black Mts, July - August, 1912, 3 males, 3 126

females (OHS, PSU). Oregon (3): Linn - Lane Co.: H J

Andrews Exp Forest, July, 1980, 3 females (OSU).

Wisconsin (2): Dane Co., June 6, 1909, male, female (UWI).

DIAGNOSIS: Coloration pale yellow to golden brown; humeri narrowly rounded, extending beyond edge of hemelytra; ventral spine not reaching hind coxae.Claspers as in fig. llc, d.

DESCRIPTION: Body length: male 8.0 - 11.0; female 8.5 -

12.0. Humeral width: male 4.1 - 5.6; female 4.4 - 6.2. Dorsum pale yellow to golden brown with brown to reddish punctation; venter pale yellow to orange-brown; punctation brown or red. Head: width: male 1.7 - 2.1; female 1.8 - 2.2; length: male 1.6 - 2.0; female 1.6 -

2.2. Antennae pale yellow to reddish orange, incisures paler. Pronotum anterolaterally concave, margined with pale yellow, dentate; median pale line extending from

calli to base. Humeri somewhat produced beyond hemelytral margins; narrowly rounded, never spinose. Scutellum: width: male 2.1 - 2.9; female 2.2 - 3.2; length: male

2.6 - 3.3; female 2.7 - 4.0. Membrane with dark band.

Legs concolorous with venter; tibiae sulcate dorsally.

Venter with row of small black dots laterally, mesally; variable in size. Ventral spine often reduced to small

protruberance not extending between hind coxae. Male

genitalia: width of genital cup 1.08 - 1.36; similar to 127

P, maculiventris; area between ventral border and ventral surface with shallow median circular depression bordered dorsolaterally by setae. Claspers triangular, concave laterally, bluntly rounded dorsal and lateral apices.

Genital plates similar to P, maculiventris;more produced ventrolaterally. Female genitalia: width of first gonocoxa 0.53 - 0.80; length 0,42 - 0.64; bulb ovate to truncate; flanges prominent; length of sclerotized rod 0.73 - 1.09.

SPECIMENS ILLUSTRATED: Humerus, genital cup: Wisconsin:

Dane Co, (UWI) (holotype); ventral spine: Wisconsin:

Dane Co. (UWI) (paratype); clasper: Montana: no locality given (MSU); spermatheca: British Columbia: Saanich District (UBC) (paratype).

DISTRIBUTION: New England and Nova Scotia west to British

Columbia, Washington, and Oregon; in the eastern U. S. south to North Carolina, Tennessee, Kentucky, northern

Louisiana, west to New Mexico and Colorado (Map 11).

MATERIAL EXAMINED: 1160 specimens from the following selected localities: CANADA: Alberta: Bilby (OSU); Edmonton (CAS, OSU, UAL, UMI); Gull Lake (UAL); High River

(OSU); Lake Wabamun (UAL); Lethbridge (UAL); Medicine Hat

(UAL); St. Paul (UAL). British Columbia: Abbotsford, Sumas Mt (UBC); Celista (UBC); Courtenay (UBC); Cowichan

Bay (CU); Golden (KU); Hagensborg (UBC); Hatzic Pr (UBC). 128

Indian River (UBC); Langley (UBC); Milner (UBC); Mission

(UBC); Nanaimo (CAS); New Westminster (UBC); Pass Creek

(UBC); Quesnel (UBC); Robson (UNC); Saanich (UBC);

Vancouver (INH, UBC); Victoria (ISU, KU, UBC); William

Lake (UBC). Manitoba: Cedar Lake (AMNH). New Brunswick:

Fredericton (UCT). Nova Scotia: 5 mi E Antigonish (OSU);

Baddeck (Cape Breton Island ) (AMNH); Boisdale (AMNH);

Grand River (CAS); Millsville (CU). Ontario: Algonquin

Prov Park (UBC); Guelph Jct (UBC); Muskoka (ISU); Ottawa

(CAS); Quetico Prov Park (DBT); Rainy River Dist (KU);

Rondeau Park (UMI); Simcoe (UCB); Swansea (ISU, LSU);

Toronto (AMNH, ISU, LSU). Quebec: Cascapedia River (UCT);

Fort Coulonge (CAS); Mt Albert, Gaspe Natl Park (AMNH);

Montreal (CAS, KU, TAM, UBC, UGA); Quinze Lake (CAS); St.

Hilaire (UBC); St. Joseph de Levis (CU).UNITED STATES:

Colorado: El Paso Co.: Manitou Springs (AMNH); Larimer

Co.: Fort Collins (CSU, KU, UAZ); Poudre River Canyon

(KU). Connecticut: Fairfield Co.: New Canaan (AMNH);

Hartford Co. (YLU); Litchfield Co.: Cornwall (UCT);

Lakeville (UMI); Litchfield (AMNH); Middlesex Co.: Nineveh Falls (YLU); Middlesex Co. (YLU); New Haven Co.:

New Haven (ISU); Tolland Co.: Mansfield Twp (UCT); Storrs

(KU, UCT). Idaho: Boise Co.: Dunnigan Creek, 13 mi E

Boise (KU); Kootenai Co.: 4 mi N Worley (UID); Latah Co.:

Moscow (UID); Moscow Mt (ISU). Illinois: Jo Daviess Co.: 129

Galena (INH). Indiana: Brown Co. (TAM); Crawford Co.:

Leavenworth (PDU). Iowa: Boone Co.: Ledges Park (ISU);

Clayton Co.: Strawberry Pt (ISU); Jackson Co.: Maquoketa

(LSU, UID); Story Co.:Ames (ISU); Tama Co.: Traer (ISU,

LSU, WSU); not located: Frazier (ISU, JE, TAM). Kentucky:

Bee Spring (AMNH). Louisiana: Winn Parish: Winnfield

(UMI). Maine: Cumberland Co.: Portland (CAS); Lincoln

Co. (OHS); Penobscot Co.: Orono (CAS); Piscataquis Co.: Baxter Park State Camp (AMNH); Baxter State Park (CU);

Sagadahoc Co.: Brunswick (AMNH); Washington Co.: Machias

(CAS); not located: Capens (CAS); Jonesboro (UAZ);

Liberty (AMNH); Maine (AMNH, CAS); Paris (CU); Township A

Range 7 (INH). Massachusetts: Barnstable Co.: Woods Hole

(AMNH); Berkshire Co.: Williamstown (AMNH); Bristol Co.:

Fall River (AMNH); Essex Co.: Beverly (AMNH); W Newberry

(YLU); Franklin Co.: Mt Toby, Sunderland (CAS); Hampden

Co.: "Fr Mills" (AMNH); Hampshire Co.: Amherst (KU);

Northampton (CAS, JE); Williamsburg (CAS); Middlesex Co.:

Framingham (AMNH, CAS); Holliston (AMNH, MCZ); Lexington

(AMNH); Mt Wachuset (AMNH); S Natick (CU); Sherborn (AMNH,

KU, SDCM); Westford (AMNH). Michigan: Alger Co.: Chatham

(MIS); Alger Co. (UMI); Allegan Co.: Fennville (MIS); Baraga Co.: L'Anse (MIS); Sturgeon River, 10 mi S L'Anse

(MIS); Baraga Co. (UMI); Barry Co. (UMI); Bay Co. (UMI);

Benzie Co.: Benzonia (UMI); Benzie Co. (UMI); Berrien Co.: E. K. Warren Preserve, Warren Woods (UMI); Charlevoix Co.: 130

Boyne Falls (UMI); (UMI); Cheboygan Co.: Burt Lake (UMI); Cheboygan (UMI); Douglas Lake (KU, UMI);

Ingleside (UMI); Cheboygan Co. (MIS, UMI); Chippewa Co.:

Drummond Island (UMI); Chippewa Co. (MIS, UMI); Clare Co.

(MIS, UMI); Clinton Co.: Bath (MIS); Crawford Co.:

Higgins Lake (UMI); Delta Co.: Escanaba (UKY); Delta Co.

(MIS, UMI); Dickinson Co. (UMI); Eaton Co.: Grand Ledge

(MIS); Olivet (UGA); Emmet Co. (UMI); Gogebic Co.: Lake

Gogebic State Park (MIS); Gogebic Co. (MIS, UMI); Gratiot Co. (UMI); Houghton Co. (UMI); Huron Co. (UMI); Ingham Co.:

E Lansing (MIS, UAZ); Ionia Co. (UMI); Tosco Co.: Oscoda

(UMI); Iron Co.: T46 R36 Sec30 (UMI); Iron Co. (UMI);

Isabella Co. (UMI); Jackson Co.: Norvell (UMI); Kalamazoo

Co.: Gull Lake Biol Stn (MIS); Kellogg Forest (UGA);

Kalkaska Co. (MIS); Kent Co.: Grand Rapids (MIS); Kent Co.

(UMI); Keweenaw Co. (UMI); Lake Co.: Big Star Lake (MIS);

Lake Co. (UMI); Lapeer Co.: Deerfield Twp (UMI); North

Branch (UMI); Luce Co.: Deer Lake State Park (MIS);

Mackinac Co.: Cedarville (CMNH); Horseshoe Bay (UMI);

Lake Michigan (UMI); Naubinway (UMI); St.Ignace (UMI);

Marquette Co.: Huron Mts (UMI); Ishpeming (WSU); Marquette

(CU, MIS); Marquette Co. (UMI); Midland Co. (MIS);

Missaukee Co. (MIS); Monroe Co.: Monroe (UMI); Montmorency Co. (UMI); Newaygo Co. (MIS, UMI); Oakland Co. (UMI);

Ogemaw Co. (UMI); Ontonagon Co.: T5ON R34W Sec8 (MIS);

Osceola Co. (MIS, UMI); Oscoda Co.: 4.4 mi S Mio (UMI); 131

Roscommon Co. (MIS, UMI); Saginaw Co. (UMI); Schoolcraft

Co.: Manistique (MIS); T42N R16W Sec13 (MIS); Schoolcraft

Co. (MIS, UMI); Shiawassee Co. (UMI); Washtenaw Co.: Ann

Arbor (SDCM, UMI); Washtenaw Co. (UAZ, UMI); Wayne Co.

(UMI); not located: Elmdale (PDU); "Ag Coil Mich" (MIS,

NDSU, UMI,UNL). Minnesota: Carlton Co.: Jay Cooke State

Park (UMN); Cass Co.: Bena (UMN); Cass Co. (UMN); Clear- water Co.: Itasca Park (UMN, WSU); Cook Co.: Lake Saganaga

(UMN); Hennepin Co.: Wayzata (UMN); Hennepin Co. (UMN);

Houston Co.: La Crescent (UMN); Winnebago Crk Valley

(UMN); Hubbard Co.: Park Rapids (LSU); Itasca Co.: T54N R26W Sec6, Clubhouse Lake (UMN); Itasca Co. (UMN); Kanabec

Co.: Mora (UMN); Lake Co.: Basswood Lake, T64N R1OW

Sec9 (UMN); Two Harbors (UMN); Norman Co. (UMN); Olmstead

Co. (UMN); Ottertail Co.: Fergus Falls (INH); Pine Co.:

Little Sand Crk, 1 mi E Cloverdale (UMN); Ramsey Co.: Mississippi River Sand Bar (UMN); Powder Plant Woods (UMN);

St. Anthony Park (UMN); White Bear (UMN); Ramsey Co. (UMN);

Rice Co: Faribault (UMN) ;St. Louis Co.: Duluth (UMN);

St. Louis Co. (UMN); Wabasha Co.: 3 mi W Wabasha (UCB); not located: Eaglesnest (INH); Fairbanks (KU); Friesland (UMN);

Kawishiwi River (UMN); Minn (UMN); Shevlin (KU). Montana:

Fergus Co.: 6 mi N Lewistown (MSU); Flathead Co.: Big

Fork (MSU); Kalispell (CAS); Gallatin Co.: Bozeman (CAS, MSU); W Gallatin Co. (MSU); Gallatin Co. (MSU); Ravalli

Co.: Hamilton (MSU). New Hampshire: Belknap Co.: 132

Barnstead (CAS); Center Harbor (CAS); Cheshire Co.:Jaf- frey (AMNH); Coos Co.: Berlin (KU); Bretton Woods (CAS,

CMNH); Dixville Notch (TAM); Mt Washington (AMNH); Randolph

(AMNH); White Mts (AMNH); Grafton Co.: Franconia (AMNH);

Hanover (CAS); Sullivan Co.: Claremont (AMNH, KU); not located: Crawfords (CAS); Glen House (CAS); Pinkham Notch

(AMNH); Three Mile Island (AMNH). New Jersey: Hunterdon

Co.: E side Clinton Reservoir (AMNH); Morris Co.: Boon- ton (UCT); Chester (UBC); Union Co.: Berkeley Heights

(AMNH); not located: Newfoundland (AMNH, KU); Potters- burg (INH). New Mexico: Colfax Co.: Thermia (UMI).

New York: Albany Co.: Rensselaerville (CU); Cattaraugus

Co.: Gowanda (CAS, KU); Chemung Co.: Elmira (UMO);

Columbia Co.: Lake Charlotte (CU); Taconic State Park

(AMNH); Delaware Co.: Cooks Falls (AMNH); Margaret (CU);

Stamford (CAS, CMNH); Erie Co.: Clarence (ISU); Hamburg

(CMNH); Lancaster (ISU); Essex Co.: Cascade Lake, Adiron- dack Mts (UCB); HeartLake (CU); Indian Falls, Mt Marcy

(CU); Keene Valley, Adirondack Mts (KU); Mt Maclntyre (CU);

Mt Marcy (CU); Mt Whiteface (CU); Franklin Co.: Saranac

Lake (CU); Genessee Co.: Batavia (UMN); Greene Co.:

Haines Falls (CU); Onteora Park (KU); Tannersville (KU);

Herkimer Co.: Little Falls (CU); Kings Co.: Brooklyn

(AMNH); Flatbush (AMNH); Monroe Co.: Rochester (AMNH);

Nassau Co.: Freeville (CU); Northwest (CU); Ontario Co.:

Stanley (CU); Orange Co.: Florida (CU); Fort Montgomery 133

(AMNH); Pine Island (AMNH); Queens Co.: Rockaway (AMNH);

Rockland Co.: Sloatsburg (AMNH); St. Lawrence Co.:

Cranberry Lake (LSU, UBC); Wanakena (AMNH); Wanakena, The

Plains (LSU); Steuben Co.: Haskinsville (CU); Suffolk

Co.: E Marion (CU); Fire Island (AMNH); Mattituck (CU);

Orient (CU); Riverhead (CU); Wyandanch (AMNH); Yaphank

(CU, KU); Tompkins Co.: Grass Bog 1 & 3 (CU); Ithaca

(AMNH, CU, UGA); McLean Bogs (CU); McLean Reservoir (CU);

Mud Creek (CU); Six Mile Creek (CU); Slaterville Spring

(CU); Snyder Heights (CU); Taughannock Falls (CU); West

Danby (CU); Van Natta's Dam (CU); Tompkins Co. (CU);

Ulster Co.: Oliverea (AMNH); Phoenicia (CAS); Slide Mt

(CU); Ulster Co. (UCT); Warren Co.: Warrensburg (KU);

Westchester Co.: Ardsley (AMNH); Pelham (AMNH); Scarsdale

(KU); White Plains (KU); not located: Lake Waccabuc (KU).

North Carolina: Caldwell Co.: Grandfather Mt (NCSU);

Haywood Co.: Blowing Rock to Linnville (AMNH); Jones Knob nr Balsam (UMI); Linnville (OSU); Linnville Falls (NCSU);

Soco Gap, Great Smoky Mts Natl Park (INH); Watauga Co.:

Blue Ridge Pkwy mi 281 (KU). North Dakota: Bottineau

Co.: Turtle Mts (NDSU); Towner Co. (DBT). Ohio: Ashland

Co. (OHS); Athens Co.: Athens (UCT); Columbiana Co.:

Salinerville (CU); Greene Co.: Clifton (OHS); Hocking Co.

(OHS, PSU, UCT); Jackson Co.: Jackson (UMN); Lucas Co.

(YLU); Richland Co.: Mansfield (AMNH); Trumbull Co. (OHS); 134

Wayne Co.: Wooster (YLU); not located "Ohio" (CU).

Oregon: Benton Co.: 4 mi NE Alsea (OSU); Alsea Mt (OSU);

Corvallis (CAS, OSU); Lobster Valley, 15 mi SW Alsea (OSU);

Marys Peak (OSU); McDonald Forest (OSU); McFadden Pond, 10

mi S Corvallis (OSU); Philomath (OSU); 3 mi NE Summit

(OSU); Clatsop Co.: Olney (CAS); Columbia Co.: Goble

(ISU); Coos Co.: Fairview, 5 mi NE Coquille (OSU);

Deschutes Co.: Sisters (WSU); Harney Co. (CAS); Hood River

Co.: Hood River (CU); Parkdale (LSU); Klamath Co.: 5 mi E Bly (AMNH); Denny Crk, Upper Klamath Lake (AMNH); Lane

H J Andrews Exp Forest (OSU); Triangle Lake (OSU);

Linn Co.: 9 mi E Holly (OSU); House Rock Cmpg (OSU);

Marion Co.: Silver Creek Falls (ISU); Washington Co.:

Forest Grove (AMNH). Pennsylvania: Allegheny Co.: Cora-

opolis (CMNH); Penn Twp (CMNH); Pittsburgh (CMNH); Bedford

Co. (CU); Butler Co.: 3 mi SE Harmony (CMNH); Centre Co.: "Bi Moshannon" (PSU); Columbia Co. (CMNH); Dauphin Co.:

Harrisburg (INH); Fayette Co.: Ohio Pyles (CMNH); Lehigh

Co.: Allentown (CU); Pike Co.: Lackawaxen (AMNH); Potter

Co.: Sweden Valley (OHS); Sullivan Co.: Laporte (PSU);

Susquehanna Co.: Dimock (MSU); Warren Co.: Youngsville

(PSU); Westmoreland Co.: Powdermill Nature Reserve (CMNH);

not located: Deer Valley (PSU); Mt Davis (PSU); "Pen Mar"

(CU); Stone Crk (PSU). South Dakota: Lawrence Co.: Lead

(ISU); Lincoln Co.: Canton (LSU). Tennessee: Blount/

Sevier Co.: Mt Leconte, Great Smoky MtsNatl Park (CU); 135

Johnson Co. 200 - 400' (OSU); Sevier Co.: Gatlinburg

(AMNH). Vermont: Caledonia Co.: St,Johnsbury (AMNH,,CAS);

Essex Co.: Gore Mt, Morton (AMNH); Lamoille Co.: Stowe

(AMNH); Orleans Co.: E Charleston (AMNH); Haystack Mt

(CAS); Washington Co.: S Barre (TJAZ); Windham Co.:

Laurel Lake, Jacksonville (UMN); Putney (AMNH). Virginia:

Grayson Co.: White Top Mt (KU); Montgomery Co.: Blacks- burg (VPI); 5 mi W Blacksburg (VPI); Nelson Co. (VPI).

Washington: Graysharbor Co.: Quinault (OHS); Jefferson

Co.: Brinnen (WSU); King Co.: Bothell (OSU); Carnation (OSU); Cedar Mt (OSU); Juanita Beach (OSU); Lake Sammamish

State Park (OSU); Maple Valley (OSU); North Bend (OSU);

Renton (OSU); Seattle (CAS, OSU, UCT); Snoqualmie Falls

(OSU); Kitsap Co.: Manchester (OSU); Pierce Co.: Electron (WSU); Elhi Hill (WSU); Fort Lewis (OSU); Puyallup (ISU,

UCB, UWI, WSU); Roy (OSU); Skagit Co.: Rockport State Park

(WSU); Samish (OSU); Snohomish Co.: Chace Lake (OSU);

Darrington (OSU); Monroe (CAS); Stillaguamish (OSU);

Sultan (OSU); Stevens Co.: Fields Springs (OHS); Thomas

Lake, 23 mi NE Colville (OSU); Thurston Co.: Millersyl- vania State Park (OSU); Olympia (OSU); Olympic Nati Forest

(OHS); Tenino (CU); Whatcom Co.: Lynden (OSU); 4 mi NE

Welcome (GMS); Whitman Co.: Pullman (WSU). West Virginia:

Barbour Co.: Philippi (CMNH); Pocahontas Co.: Cranberry

Glades (KU); Cheat Mts (CMNH).Wisconsin: Bayfield Co.: Lake Namekagon (UWI); Bayfield Co. (UWI); Clark Co. (UWI); 136

Columbia Co. (UWI); Dane Co.: Madison (UWI); UWI Campus

(UWI); Dane Co. (UWI); Dodge Co.: Beaver Dam (CAS); Door

Co.: Washington Island (UWI); Door Co. (ISU, UWI);

Florence Co. (UWI); Fond du Lac Co. (UMI); Grant Co.

(UWI); Jackson Co. (UWI); Jefferson Co. (UWI); La Crosse

Co. (UWI); Marinette Co. (UWI); Oconto Co.: Lakewood

(MIS); Oneida Co. (UWI); Polk Co.: Gibson Lake (UWI);

Polk Co. (UCB); Price Co.: Phillips (UWI); Richland Co.

(UWI); Sauk Co.: 3 mi SE Sauk City (UGA); 10 mi W Sauk

City (UGA); Trempealeau Co. (UWI); Vernon Co.: Vernonia

(UWI); Vernon Co. (UWI); Walworth Co.: Lake Geneva (UWI); not located: Leroy (MIS).

This species has long been known under the name P. modestus (Dallas). I examined the type specimen of Arma modesta Dallas and found the male claspers, ventral spine, and body size to be indistinguishable from P. maculiventris. I therefore synonymized the 2 species.

also examined the type of Rhaphigaster aggressor Walker,

1867, listed by Kirkaldy (1909) as a synonym of P. modestus, and in the collection of the British Museum

(Natural History). This specimen, collected at Santo Domingo, Dominican Republic (Hispaniola) is not related

to either P. modestus (=maculiventris) or P. brevispinus,

as it is a Neotropical form, probably P. jole

American workers have differentiated between P.

maculiventris and P. brevispinus (heretofore known as P. 137 modestus) based on its short ventral spine and smaller size (McPherson 1982, references therein). It may be separated from P. serieventris by its lighter, unmottled coloration and clasper characteristics (fig. lic, d).The spermathecal bulb of brevispinus is shorter than that of maculiventris or serieventris and is usually truncate.

P. brevispinus is rather constant morphologically over its range. Minor differences in the degree of concavity of the lateral edge of the clasper exist. The ventral spine may be a small, barely perceptible swelling or a more defined, pointed structure as in fig. llf. Large females of darker than average coloration may be confused with P. serieventris but the former's more narrowly rounded humeri and shorter, truncate spermathecal bulb separate the two.

All biological information concerning P. brevispinus is listed under P. modestus (Dallas) because of the former's association with a type specimen that is a synonym of P. maculiventris. Eggs and life history have been described (Coppel and Jones 1962; Tostowaryk 1971).

Results are given in Table 1. Tostowaryk (1971), in the most comprehensive study to date, described this species as an abundant predator of sawfly larvae (Neodiprion spp.) in jack-pine stands and understory in Quebec. Hunting, feeding, and mating behavior were observed. Tostowaryk (1971) found P. brevispinus to be a generalist predator in food preference tests. Studies of the functional 138 response of this species showed that as prey density rose, predator success did so at a decelerating rate (Tostowaryk

1972). The author attributed this to increased prey defense, as predators attacking groups of sawfly larvae received a coating of sticky regurgitate which took time to remove from their tarsi, antennae, and rostrum. Kirkland (1898) observed P. brevispinus more often on bushes than on trees in Massachusetts. Blatchley (1926) mentions its occurrence on hazel. I have taken it in small numbers on oak, red alder, and willow in Oregon.

McPherson (1982) lists 48 insect species upon which

P. brevispinus is known to prey. Additional prey species are shown in Table 2. More information about the habits and potential use of P. brevispinus in integrated pest management programs for forest and ornamental pests is needed.

Podisus fretus Olsen (Fig. 12; Map 12)

Podisus fretus Olsen 1916: 82; Van Duzee 1917: 79.

TYPE AND TYPE LOCALITY: Podisus fretus Olsen, holotype, male, fair condition (AMNH): "Rockaway Beach, L. I. /

4 - 17 - 1912 / Coll. by A. Nicolay / TYPE (red) / C. E.

Olsen Collection / Podisus fretus; Chris. E. Olsen /

Holotype; Podisus fretus; C. E. Olsen (red)." Allotype, 139 female, poor condition (AMNH): "Rockaway Beach; V - 7 -

10 L. I. / Coll. Chris. E. Olsen / Allotype (red) / C. E.

Olsen Collection / Podisus fretus; Chris. E. Olsen /

Allotype; Podisus fretus; C. E. Olsen (red)." Paratypes

(AMNH), male, fair condition: "Wyandanch, L. I.; 8 - 30 -

1914; F. M. Schott / Paratype (red) / C. E. Olsen

Collection / Podisus fretus; Chris. E. Olsen;" female, fair to good condition: "Wyandanch, L. I.; 8 - 30 - 1914

F. M. Schott / Paratype / Podisus fretus Chris. E. Olsen."

Metatype (AMNH), male, fair - good condition: "Wyandanch,

L. I.; Sep. 17, 1916; F. M. Schott / Metatype (red) /

C. E. Olsen Collection / Apateticus (Podisus) fretus

Olsen."

DIAGNOSIS: Length 10 - 14 mm; coloration reddish - brown; humeri bluntly rounded; venter with irregular reddish ventral spots. Claspers as in figure 12c, d.

DESCRIPTION: Body length: male 10.0 - 13.5; female

12.5 - 14.0. Humeral width: male 5.8 - 7.0; female 6.6 -

7.3. Dorsum light brownish yellow with dense red punctation lending ruddy brown appearance; venter pale to orange yellow with profuse red punctation. Head thickly punctured with reddish brown; width: male 2.4 -

2.6; female 2.5 - 2.8; length: male 2.1 - 2.4; female

2.2 - 2.5. Antennae reddish brown, particularly segments

III - IV; incisures paler. Pronotum with straight to 140 slightly concave anterolateral margins; explanate and dentate on anterior half. Humeri blunt, rounded, not much produced beyond edge of hemelytra. Scutellum: width: male 3.0 - 3.7; female 3.4 - 3.9; length: male 3.4 - 4.9; female 4.1 - 5.1. Hemelytra each with irregular brown spot near apex. Membrane with dark band. Legs ochraceous to rufous; punctate with red or brown; tibiae of some specimens bright red.Venter with series of irregularly- shaped reddish brown spots mesally.. Ventral spine extending between hind coxae. Male genitalia: width of genital cup 1.66 - 1.84; lateral lobes rounded, margins deflexed. Claspers with wide triangular dorsal 2/3 - 3/4; ventrally narrow, tapered to rounded lateral apex.

Genital plates subquadrate, dentate; mesal and dorsal margins rounded; lateral border scalloped; surface with flattened tubercles and ridges. Female genitalia: first gonocoxae deflexed and flattened; inner posterior margins with excavated appearance; this area with brown spot; width 0.74 - 0.80; length 0.64 - 0.80. Bulb elongate, constricted near tip; flanges small; length of scierotized rod 1.06 - 1.40.

SPECIMENS ILLUSTRATED: Humerus: Florida: Taylor Co.

(UMI); clasper: Florida: Taylor Co. (UMI); spermatheca:

Florida: Taylor Co. (UMI). 141

DISTRIBUTION: Massachusetts to Florida, Indiana, and

Michigan (Map 12).

MATERIAL EXAMINED: 25 specimens from the following localities: Connecticut: Windham Co.: Putnam (AMNH).

District of Columbia: (CU). Florida: Taylor Co. (UMI).

Georgia: Clarke Co.: 3 mi N Athens (OSU). Indiana:

Lake Co. (PDU). Massachusetts: Barnstable Co.: Orleans

(AMNH, CAS); Hampshire Co.: Amherst (UMO). New Jersey:

Middlesex Co.: New Brunswick (AMNH); Ocean Co.:

Lakehurst (CU). New York: Queens Co.: Rockaway (AMNH);

Suffolk Co.: Wyandanch (AMNH). North Carolina: "N. C."

(ISU). South Carolina: Georgetown Co.: Huntington Beach

State Park (UKY). Virginia: King & Queen Co. (VPI).

This species is the largest Nearctic Podisus. It is rarely collected and of limited distribution. Olsen (1923) believed that P. fretus might be overlooked because of its resemblance to more common species such as P. maculiventris and P. serieventris. The combination of large size, blunt humeri, long ventral spine, and ruddy coloration suffices to distinguish fretus from its congeners.

The claspers are distinctive in their broadly triangular upper 1/2 to 3/4 (fig. 12c). The ventral part of the clasper is narrower proportionately than in other

Podisus. Clasper shape varies in the steepness of the lateral and mesal edges and breadth of the dorsal portion. 142

Nothing is known of the immature stages, life cycle, or habits of this uncommon species. Hetrick (1959) recorded it as a predator of the blackheaded pine sawfly,

Neodiprion excitans Rohwer, in an outbreak on loblolly pine in north central Florida. Olsen (1916) mentioned P. fretus' occurrence in beach wash-up on Long Island as well as on oak and pine.

Podisus placidus Uhler

(Fig. 13; Map 13)

Podisus placidus Uhler 1870: 203, fig. 124a, b; 1897b: 116;

Kirkland 1898: 416; Van Duzee 1904: 71.

Apateticus (Eupodisus) placidus Schouteden 1907: 72.

Apateticus (Podisus) placidus Kirkaldy 1909: 20.

TYPE AND TYPE LOCALITY: Podisus placidus Uhler,

"lectotype" (no red label), female, very good condition

(USNM): "Maiden, Mass. / P. R. Uhler Collection / Podisus placidus Uhl.; See Type Collection. ("See," is written over with black ink to read, "Lec," and "Collection" is crossed out making the label appear to read, "Lec Type

Gelleetien.") I will propose that this specimen be given lectotype status in a future publication. The type locality is Malden, Massachusetts. 143

DIAGNOSIS: Anterolateral pronotal margins straight; humeri blunt; hemelytral membrane without dark band.

Claspers and spermathecal bulb as in figure 13c, d, e.

DESCRIPTION: Body length: male 8.0 - 10.0; female 9.0 -

12.0. Humeral width: male 4.3 - 5.3; female 4.8 - 6.2. Dorsum light yellow to orange brown, some specimens fuscous or piceous; punctation red or brown; venter pale yellow to reddish orange; red or brown punctate. Head finely outlined with black; width: male 1.9 - 2.1; female 2.0 -

2.4; length: male 1.6 - 1.9; female 1.7 - 2.1. Antennae yellow to brown; piceous in dark specimens; second segment often darkest. Pronotum with straight anterolateral margins outlined in pale yellow (orange in dark specimens); anterior 1/3 to 1/2 denticulate. Humeri bluntly rounded, not produced. Scutellum: width: male 2.4 - 3.0; female

2.6 - 3.4; length: male 2.7 - 3.6; female 3.1 - 4.1.

Membrane golden, lacking dark band. Legs yellow or orange to piceous, thinly setose; tibiae sulcate dorsally.

Ventral spine slender, reaching between hind coxae. Male genitalia: width of genital cup 1.22 - 1.49; lateral lobes produced, tapered apically; area between ventral border and lower edge of cup concave; claspers boot-shaped; thin straight process extends dorsad above gradually tapered laterally directed "foot;" latter with small subapical tooth. Genital plates quadrate, rounded mesally; strongly indented laterally; dorsal margin 144 deflexed, surface ridged or with deflexed tubercles.

Female genitalia: first gonocoxa: width: 0.52 - 0.80; length 0.48 - 0.66. Bulb round with small flanges; length of scierotized rod 1.05 - 1.62.

SPECIMENS ILLUSTRATED: Humerus: New York: St. Lawrence

Co.: Wanakena (AMNH); genital cup: New York: Tompkins

Co.: Ithaca (CU); clasper: CANADA: Manitoba: Mafeking

(KU); spermatheca: Oregon: Wasco Co.: T2N R12E Sec5

(GMS).

DISTRIBUTION: Across the northern United States and Canada south to Oregon, UtahColorado, Oklahoma, Louisiana,

Illinois, Indiana, Ohio, and Pennsylvania (Map 13).

MATERIAL EXAMINED: 676 specimens from the following localities: CANADA: British Columbia: Agassiz (INH,

UBC); Goldstream (UBC); Robson (UBC); Saanich Dist (UBC);

Univ British Columbia (UBC). Manitoba: Aweme (UBC); 16 mi E Aweme (UAL); Hartney (KU); Mafeking (KU). Ontario:

Ottawa (CAS); Toronto (LSU). Quebec: Montreal (JE);

Montreal, Beaulieu (CAS); Ormstown (JE). UNITED STATES:

Arkansas: Washington/Crawford Co.: Devil's Den State

Park nr Winslow (UMO). Colorado: Boulder Co.: Berkley

(OSU); Co.: Denver (AMNH, CAS, CU); El Paso Co.:

Colorado Springs (CAS); Rock Creek, vic Colorado Springs

(AMNH); Jefferson Co.: Golden (UMI); Morrison (KU);

Larimer Co.: Fort Collins (CAS, CSU, KU, PSU, UAZ); 145

Morgan Co,: Snyder (CSU, KU). Connecticut: Fairfield

Co.: New Canaan (ANMH); Stamford (AMNH); Westport (AMNH);

Litchfield Co.: Lakeville (UMI); Litchfield (AMNH);

Litchfield, Mohawk Mt (AMNH); Pleasant Valley (KU); West

Cornwall (AMNH); Middlesex Co.: Cromwell (CAS); Portland

(YLU); New Haven Co.: 1 mi SSW Bethany Ctr (YLU); Meriden

(UCT); New Haven (CU, ISU, UCT, YLU); North Haven (KU);

S Meriden (KU, YLU); New London Co.: N Stonington (CAS);

Stonington (YLU); Tolland Co.: Mansfield (UCT); Mansfield

Twp, Storrs (UCT); Stafford Twp nr Crystal Lake (UCT);

Storrs (OSU, UCT); Windham Co.: Putnam (AMNH); Willimantic

(UCT); not located: Bozeah Twp (UCT); Colebrook (CAS);

Savin Rock (YLU). Idaho: Jefferson Co.: Rigby (ISU);

Madison Co.: Rexburg (UID); not located: Long (KU).

Illinois: Champaign Co.: Champaign (KU); Jersey Co.:

Principia College (YLU); Henderson Co.: Henderson State

Forest, Oquawka (UMN); Johnson Co. (INH); Mason Co.:

Havana, Devil's Hole (INH); Mason State Forest (AMNH);

Mason Co. (UCR). Indiana: not located: Mineral Springs

(OSU). Iowa: Boone Co.: Ledges State Park (ISU, TAM);

Cass Co.: Atlantic (TAM); Linn Co,: Expt Stn, Mt Vernon

(ISU); Story Co.: Ames (ISU, UAZ, UWI); Expt Stn Ames

(ISU); Tama Co.: Traer (ISU, LSU); Van Buren Co.:

Keosauqua (ISU); Washington Co.: Crawfordsville (ISU) ;

Woodbury Co.: Sioux City (UMN). Kansas: Leavenworth Co.:

Fort Leavenworth (UMO). Louisiana: Winn Parish: Winnfield 146

(UMI). Maine: Cumberland Co.: Gorham (MIS); Piscataquis

Co.: Mt Katandin (AMNH); not located: Paris (CU).

Massachusetts: Barnstable Co.: Elizabeth Island (MCZ);

Penekise Island (MCZ); Provincetown (AMNH); Woods Hole

(AMNH); Berkshire Co.: Bird Sanct, Lenox (KU); Bristol

Co.: Taunton (AMNH); Essex Co.: Saugus (CAS); Hampshire

Co.: Amherst (MIS, MSU, UGA, UMO); Northampton (JE);

Hampden Co.: Chicopee (CU); Middlesex Co.: Framingham

(AMNH, SDCM); Holliston (AMNH, MCZ); Lexington (AMNH);

Lincoln (AMNH); Littleton (UMI); Malden (AMNH, CU, KU); Natick (CAS, SDCM); Newton (CAS); Sherborn (CAS, SDCM);

Sudbury (KU); Norfolk Co.: Milton (CMNH, UMI); Wellesley

(AMNH CAS); Suffolk Co.: Boston (CAS); Cambridge (AMNH);

Worcester Co.: Ashburnham (OSU); not located: Blue Hills

(CAS); Forest Hills (CAS); Silver Hill (UCR). Michigan:

Alcona Co. (OHS, UMI); Allegan Co.: Allegan (MIS);

Allegan State Forest (MIS); Alpena Co. (MIS); Barry Co.:

T3N RlOW Sec26 (MIS); Yankee Spring Rec Area (UCB); Yankee

Springs (MIS); Orangeville (MIS); Charlevoix Co.: Boyne

City (UMI); Norwood Twp (UMI); Cheboygan Co.: Cheboygan (UMI); Douglas Lake (KU, UMI); Ingleside (UMI); Cheboygan

Co. (KU, MIS, OSU, UMI); Clare Co.: T17N R6W SEc18 (UMI);

Clinton Co. (MIS); Crawford Co.: T26N R2W Sec33 (UMI);

Crawford Co. (UMI); Grand Traverse Co.: Traverse City

(MIS); Gratiot Co. (UMI); Huron Co.: Sand Point (UMI);

Ingham Co,: Aurelius (MIS); Jackson Co.: Waterloo (DBT); 147

Kalamazoo Co.: Gull Lake Biol Stn (MIS); Kalkaska Co.

(UMI); Lake Co. (UMI); Lapeer Co. (UMI); Livingston Co.:

E. S. George Reserve (UMI); Livingston Co. (UMI); Mackinac

Co.: St. Ignace (UMI); Manistee Co.: Dublin (MIS);

Manistee Co. (UMI); Midland Co. (ISU, MIS); Montcalm Co.

(UMI); Montmorency Co.: Atlanta (MIS); Newaygo Co.:

Newaygo (UMI); White Cloud (MI); Oakland Co.: Cooper

Woods (UMI); Farmington (UMI); Milford (UMI); New Hudson

(UMI); Oakland Co. (UAZ, UMI); Oscoda Co.: Luzerne (MIS);

Oscoda Co. (UMI); Otsego Co,: Vanderbilt (MIS); Otsego

Co. (UMI); Roscommon Co.: Higgins Lake (UMI); Tuscola Co.

(UMI); Van Buren Co.: Grand Junction (MIS); Washtenaw Co.: Ann Arbor (UMI); Ann Arbor, 3rd Woods (UMI); 3rd Sister

(UMI); Washtenaw Co. (KU, UMI); Wayne Co.: Detroit (UMI);

Wexford Co.: Cadillac (MIS. UMN); Mesich (MIS); Woodward

Lake (MIS); Wexford Co. (UMI); not located: "Ag Coll Mich"

(UMI); "Mich" (AMNH, CSU). Minnesota: Carlton Co.:

Cloquet (UMN); Cass Co. (UMN); Clearwater Co.: Itasca

Park (WSU); Lake Itasca (DBT); Hennepin Co. (UMN); Mille

Lacs Co. (UMN); Olmstead Co.: Rochester (UMN); Olmstead

Co. (UMN); Ottertail Co.: Fergus Falls (INH); Pine Co.

(UMN); Ramsey Co.: St. Anthony Park (UMN); Ramsey Co.

(UMN); St. Louis Co.: Duluth (INH, UMN); St. Louis Co.

(UMN); Wabash Co.: Lake City (UMN). Missouri: Boone Co.:

Ashland (UMO); Ashland Wildlife Area (UMO); Iron Co. (UMO);

Newton Co.: Stark City (UMO); Phelps Co.: St. James 148

(UMO); Phelps Co. (UMO); St. Louis Co.: Ranken (UMO).

Montana: Flathead Co.: La Salle (UMI); Gallatin Co.

(MSU); Missoula Co.: Missoula (MSU); Ravalli Co.: Como

(MSU); Teton Co.: Choteau (MSU). Nebraska: Cass Co.:

South Bend (UNL); Dakota Co.: S Sioux City (UNL); Sioux

Co.: Monroe Canyon (UNL); Sioux Co. (UML); not located:

War Bonnet Canyon (UNL). New Hampshire: Belknap Co.:

Barnstead (CAS); Conway (MIS); Cheshire Co.: Jaffrey

(UMI); Strafford Co.: Dover (AMNH, OSU); New Durham

(AMNH). New Jersey: Bergen Co.: Bear Swamp nr Ramsey (AMNH); Bear Swamp, Ramapo Mts (UCB); Mahwah (UGA); Ramsey

(AMNH); Monmouth Co.: 3 mi SE Farmingdale (UMI); Morris

Co.: Morristown (JE); Ocean Co.: Holgate Unit of B. N.

W. R., Long Beach Island (INH); Lakehurst (UCT); Long

Beach Island (AMNH); Passaic Co.: Passaic Jct (AMNH);

Somerset Co.: Bernardsville (UCT); Sussex Co. (CU); not located: Glen Cove (UCT); Newfoundland (AMNH). New York:

Allegany Co.: Cuba (MIS); Chemung Co. (UMO); Delaware

Co.: Margaretsville (CU); Essex Co.: Mt Maclntyre (CU); Westport (AMNH); Greene Co.: Onteora Park, 2400 - 2600',

Tannersville (KU); Kings Co.: Flushing (AMNH); Monroe

Co.: Rochester (AMNH); Rochester Jct (AMNH); Orange Co.:

Fort Montgomery (KU); Pine Island (AMNH); Queens Co.:Far

Rockaway (AMNH); Rockaway Beach (KU); St. Lawrence Co.:

Parishville (UMI); Wanakena (AMNH); Suffolk Co.: Babylon

(CU); Central Port (AMNH); Coram (AMNH); East Hampton 149

(KU); Flanders (CU); Greenport (CU); Huntington (AMNH);

Mastic (AMNH); Montauk (AMNH); Oakdale (AMNH); Orient

(CU); Pinelawn (AMNH); Riverhead (PDU); Sag Harbor (CU);

Southold (CU); Wading River (AMNH); Wyandanch (AMNH);

Yaphank (AMNH, KU, PDU); Tompkins Co.: Ithaca (AMNH, CU,

PDU); Sphaerium Brook, McLean Res (CU); Six Mile Creek

Valley (CU); Tompkins Co, (CU); Ulster Co. (UCT); Warren

Co.: Lake George (AMNH); Westchester Co.: Ardsley (AMNH);

Bronxville (AMNH); Lewisboro (AMNH); Scarsdale (KU);

Westchester Co. (OSU); Wyoming Co.: La Grange (DBT, UMO); not located: Aqueduct (AMNH); Bald Hill, Caroline (CU);

Hunters Island (LSU); Karner (CMNH). North Dakota:

Billings Co.: Theodore Roosevelt Memorial Park (NDSU);

Cass Co.: Fargo (MDSU); Valley Co.: Beach (NDSU). Ohio:

Athens Co.: Athens (UCT); Delaware Co. (OHS); Fulton Co.

(YLU); Portage Co. (YLU). Oklahoma: Payne Co.:

Stillwater (OKS); not located: Briartown (UMO). 21-222a:

Linn Co. (OSU); Wasco Co.: R12E T2N Sec5 (GMS); R11E T2N

Seci (GMS). Pennsylvania: Allegheny Co.: Pittsburgh

(CMNH); Cambria Co.: Johnstown (CSU); Centre Co.: State

College (PSU); Franklin Co.: Horse Valley (PSU); Monroe

Co.: Delaware Water Gap (AMNH); Stroudsburg (UCT);

Sullivan Co.: Laporte (PSU); Warren Co.: Youngsville

(PSU); Washington Co.: Canonsburg, Mt Blain (UMI); Oakmont

(OHS); Washington Co. (CMNH); not located: Charter Oak

(PSU); Eastern PA (UWI); Mt Alto (PSU). South Dakota: 150

Butte Co.: Newell (LSU); Nisland (LSU); Lawrence Co.:

Cheyenne Crossing, Black Hills Natl Forest (MIS); Lead

(ISU); Roberts Co.: Sissiton (ISU). Utah: Cache Co.:

Brigham (OSU); Logan (LSU); Davis Co.: King's Stn (CAS);

Salt Lake Co.: Salt Lake City (CAS, KU, OSU, UCB); Utah

Co.: American Forks Canyon (CAS); Weber Co.: Ogden

(KU). Vermont: Franklin/Lamoille Co.: Green Mts (CSU).

Washington: King Co.: Seattle (OSU); Snoqualmie Falls

(OSU); Mason Co.: Shelton (OSU); Stevens Co.: Pleasant

Valley (UID); Yakima Co.: N Yakima (LSU). Wisconsin:

Bayfield Co.: Bayfield (ISU, UWI); Burnett Co.: N of

Riverside (UWI); Columbia Co. (UWI); Crawford Co.: Gays

Mills (UWI); Wauzeka (UWI); Dane Co.: Madison (UWI);

Dane Co. (UWI); Door Co. (UWI); Douglas Co.: Lake

Nebagamon (UCR); Dunn Co. (UWI); Green Lake Co. (UWI);

Jackson Co. (UWI); Juneau Co. (UWI); La Crosse Co. (UWI);

Marathon Co.: Rib Mt State Park (KU, UMN); Oconto Co.:

Lakewood (MIS); Pierce Co. (UWI); Polk Co.: Gibson Lake

T34N R16W Sec34 (UWI); Portage Co.: 2.2 mi E Plover

(UWI); Sauk Co.: Castle Rock (UGA); Partray's Glen

(UMO); Sauk City (UGA); Trempealeau Co. (UWI); Vilas Co.

T42N R8E (UWI); Vilas Co, (UWI); Waukesha Co. (UWI); Wood

Co.: Griffith State Nursery (UWI); Nepeo Lake, Nepeo

Boys Camp (UWI). Wyoming: Platte Co.: Wheatland T25N R67W Sec22 (UWY). 151

Saunders (1870) reported on a "friendly bug" preying on larvae of the imported currantworm, Nematus ribesii

(Scopoli) which a professor Ulke of Baltimore, Maryland named Arma placidum (sic). No written description was published under this name, although Saunders (1870) did provide an illustration of the insect.Kirkland's

(1897) comments indicate that specimens were sent to

Uhler, not Ulke as Saunders (1870) indicated. Uhler's published descriptions (1870, 1897b) give the name as

Podisus placidus which is listed first in the synonymical bibliography.

P. placidus varies in coloration in some parts of its range. I examined specimens from Washington that were piceous with orange anterolateral pronotal margins. The male and female genitalia of these individuals were indistinguishable from lighter-colored specimens.

The male claspers (fig. 13c, d) vary in the steepness of the curve between the dorsal and ventral portions. The subapical tooth may be rounded or prominent and acute.

Females show little variation in the characteristically round spermathecal bulb. Kirkland (1898), Coppel and Jones (1962), and Oetting and Yonke (1971) studied the life cycle of this species.

Results are summarized in Table 1. Nymphal instars were described by Oetting and Yonke (1971) who observed aggregation of first instars and reported that nymphs were 152 aggressive predators.

Warren and Bjegovic (1972) noted differences in responses to photoperiod by P. placidus and P. maculiventris. The former went into reproductive diapause at 9 and 12 hour photoperiods while the latter was active under light regimes of 9, 12, 14, and 16h at 80°F. All of these studies reported that P. placidus was a generalist predator. A list of known prey is given in Table 2.

Telenomus persimilis Ashmead (Hymenoptera:

Scelionidae) parasitizes the eggs of P. placidus (Table 3).

Podisus saqitta (Fabricius)

(Fig. 14; Map 14)

Cimex saqitta Fabricius 1794: 99.

Pentatoma didyma Palisot de Beauvois 1805: 128.

Arma didyma Amyot & Serville 1843: 85.

Telepta didyma Stal 1862: 90.

Podisus didyma Stal 1867: 497.

Arma monospila Walker 1867: 136.

Podisus saqitta Distant 1880: 37, pl. 1 fig. 22; Van Duzee

1904: 72.

Podisus (Podisus) saqitta S.W. 1870: 50; Van Duzee 1917:

80.

Apateticus (Eupodisus) saqitta Schouteden 1907: 72. Apateticus (Podisus) saaitta Kirkaldy 1909: 20. 153

TYPES AND TYPE LOCALITIES: Cimex sagitta Fabricius,

probably destroyed (see Zimsen 1964). Pentatoma didyma Palisot de Beauvois was notseen by the end of this study,

but I expect to see itsoon. Arma monospila Walker, type, female, fair condition (BM): "Type (circled) / 66; 12- /

16. Arma monospila." Fabricius' type locality is

"American meridionalis insulis." Walker's specimen was collected in Mexico. I have so far been unable to obtain a copy of Palisot de Beauvois' original description of

Pentatoma didyma and therefore cannot accurately state the type locality.

DIAGNOSIS: Humeral spines notched apically; tipped with

rusty red to fuscous. Claspers as in figure 14c, d. Spermathecal bulb fusiform.

DESCRIPTION: Body length: male 10.0 - 13.0; female 12.0

13.5. Humeral width: male 5.8 - 7.5; female 6.6 - 8.3.

Dorsum golden brown, at times with light gray wash; punctation light red- brown; venter pale yellow to yellowish brown with brown or red punctures. Head thinly margined with fuscous; width: male 2.0 - 2.4; female 2.2 - 2.5; length: male 1.7 - 2.1; female 1.8

2.3. Antennae pale to brownish yellow, segments IV and V often darker. Pronotum with concave, pale, dentate anterolateral margins. Humeri spinose; spines emarginate reddish to fuscous, often curved forward. Scutellum: 154 width: male 2.8 - 3.4; female 2.7 - 3.7; length: male

3.3 - 4.1; female 3.8 - 4.7.Membrane golden with dark apical band. Legs pale yellow to yellowish brown; tibiae sulcate dorsally. Venter with black dots laterally; sternite VI often with large, black spot mesally.Ventral spine extending between hind coxae. Male genitalia: width of genital cup 1.40 - 1.50; cup very similar to P. maculiventris and P. serieventris; area between ventral border and lower surface of cup shallowly concave with small median elevation. Claspers with lateral edge deeply concave; lower portion broadly rounded at lateral apex.

Genital plates subquadrate; mediodorsal edge rounded, irregular; lateroventral portion slightly extended; surface with incomplete transverse ridges. Female genitalia: width of first gonocoxa 0.58 - 0.80; length 0.53 - 0.66.

Bulb spindle-shaped; flanges small, thickened; length of sclerotized rod 0.75 0 1.28.

SPECIMENS ILLUSTRATED: Humerus, genital cup: Texas:

Cameron Co.: Brownsville (UCT); clasper: Texas: Cameron

Co.: Sabal Palm Grove Sanctuary (UGA); spermatheca:

Texas: Hidalgo Co. (OHS).

DISTRIBUTION: Southeastern Texas, southern Florida,

Mexico, Guatemala, Honduras, Costa Rica, Nicaragua, and the Caribbean (Map 14). 155

MATERIAL EXAMINED: 145 specimens from the following localities: COSTA RICA: Cartago: Pacayas (AMNH).

Guanacaste: Comelco, 4 km NW Bagaces, Area 2 (UCB).

San Jose: San Jose (TAM, UCB). CUBA: Havana (KU);

Soledad (AMNH); N of Venales (AMNH). DOMINICA: Long

Ditton (AMNH). DOMINICAN REPUBLIC: S4nchez (AMNH).

GUATEMALA: Peten: Poptun (UMI); Tikal (UMI).

Sacatepeguez: Amatitlan (AMNH); Antigua (AMNH, ISU, LSU);

Capetillo (AMNH); Reunidn (AMNH). Suchitepequez:

Variedades (AMNH). Zacapa: Zacapa (AMNH). Not located: Duenas (AMNH); Finca Barcona (AMNH); "Guatem." (AMNH).

HAITI: Port au Prince (UAZ). HONDURAS: Atlantida: La

Ceiba (AMNH). MorazAn: Escuela Agricultura Panamericana Zamorana (UMI); Lancetilla (UMI). JAMAICA: Halishire Hills (DBT); Hardwar (AMNH); Palm Beach, Montego Bay (KU).

MEXICO: Chiapas: 4 mi W San Crist6bal (UMI); Santo Domingo, 15 mi SW Simojovel (UCB); Suchiapa (UCB);

Tapachula (DBT); Tonal (AMNH). Distrito Federal:

Amecameca (PSU); Chalco (PSU). Guerrero: 19 mi S Iguala

(TAM); 3.7 mi E Marquelia (TAM). Jalisco: Ajijic (UAZ); 4 mi W Mazamitla (AMNH); NW Slope Nevado de Colima, El

Isolte 7500' (UMI). Mexico: Real de Arriba, Temescaltepec

(UCB); Tejupilco, Temescaltepec (UCB). Michoacan: btwn

San Juan Par & Zirosto (UMI). Morelos: Cuernavaca (CAS,

PSU); 13 km E Cuernavaca (UCB). Nayarit: Arroyo de

Santiago nr Jeslls Marfa (UCB); Tepic (CAS). Oaxaca: 156

27 mi E Juchitan (TAM); 8 mi N La Ventosa (UCB); Oaxaca

(AMNH); 13 mi S Suchixtepec (TAM); 3 mi N Tapanatepec

(TAM). Puebla: Matamoros (UMI); Puebla (UCB). San Luis

Potosi: El Salto (KU); Tamazunchale (CAS, DBT); WSW

Valles (Covadonga) (AMNH). Tamaulipas: Bocatoma, 6 mi S

Gdmez Farfas (CBT); Ciudad Victoria (ISU); Grutas de

Quintero, 8 mi S Ciudad Mante (UGA); 17 mi N Victoria, Rio

Santa Engracia (UMI). Veracruz: Cotaxtia Exp Stn, Cotaxtla (UCB); Jicaltepec (ISU, OHS); Orizaba (UMI); San

Rafael Jicaltepec (ISU, OHS). Yucatan: Colonia Yucatan

(AMNH). Not located: "Atlantic Viscount, Mex" (CSU).

NICARAGUA: Lean: Hacienda Casa Blanca (JE). PUERTO RICO:

Adjuntas (AMNH); Fortuna A. E. S. (OHS); Jayuya (AMNH); Loquillo Natl Forest (AMNH); Mayaguez (TAM); San German

(TAM); Tallaboa nr Ponce (AMNH). UNITED STATES: Florida:

Broward Co.: Hallendale (AMNH); Dade Co.: Homestead

(OSU); Palm Beach Co.:Royal Palm Park (PDU); not located: Princeton (UCT).

Podisus fuscescens (Dallas) might be confused with P. saqitta. Van Duzee (1917) listed the former as occurring north of Mexico, but it is a Neotropical species. I examined the type specimen of Arma fuscescens Dallas, female, poor condition (BM): "Type (circled) / Mexico /

(white rectangle with a handwritten letter "a" written sideways) / 7. Arma fuscescens." This species is darker brown dorsally than saqitta, brown ventrally, unlike 157

saqitta which is pale yellow, and has straight humeral

spines that are only slightly notched. I examined other

specimens of fuscescens (AMNH) and found the claspers to be very different from saqitta. The spermathecal bulb is elliptical rather than fusiform as in sagitta. Most of P. saqitta's distribution is Neotropical.

Its United States distribution includes an area in southern Texas corresponding to Bailey's (1978) Mesquite

Acacia Savanna Section and the Florida Everglades. No habitat or host plant data were available.

This species was the most abundant pentatomid predator on the Mexican bean beetle (Epilachna varivestis) at Cuernavaca, Morelos, Mexico during August and September

(Plummer and Landis 1932). The authors reared P. saqitta in the laboratory and obtained life history data which are presented in Table 1. Egg masses contained 20 - 50 eggs and the development period was 62 - 64 days. Plummer and Landis (1932) believed this to account for the species' scarcity until bean beetle populations passed their peaks.

The authors felt that P. saqitta was a valuable predator with an average daily food equivalent of 3.5 third instar beetle larvae.

Painter (1955) collected P. saqitta on corn in a study of corn and teosinte insects in Guatemala. Stoner

(1922) recorded the species from Barbados. In Mexico,

Gonzdlez (1978) recorded Trissolcus sp. as an apparently 158 host-specific egg parasitoid of P. saqitta (Table 3). 159

CONCLUSIONS

Apateticus and Podisus should be maintained as

separate genera based on genitalic and external morphological characters. The former consists of 5

species in 2 subgenera, all occurring in the Nearctic

region. Podisus comprises 8 Nearctic species. P. concirex,

although Neotropical, is included as an incidental

Nearctic species. P. fuscescens, listed by Van Duzee

(1917) as occurring in the United States, is a Neotropical

species. Podisus brevispinus, heretofore known as P. modestus because of its association with a type specimen which is actually synonymous with P. maculiventris, is

described as a new species.

The species are differentiated on the basis of morphological characters. The shape of the male claspers will distinguish all but 2 pairs of species, A. marginiventris - A. lineolatus and P. maculiventris-

P. serieventris. Pronotal shape, color, and length of the ventral spine will separate each of these pairs. Female genitalia are not reliable for species-level discrimination because of their uniformity. A few species have

characteristic spermathecal bulbs, but when the entire genus Podisus is examined, the taxonomic value of these 160

distinctions could diminish.

Biological studies show these insects to be

generalist predators. P. maculiventris, P. serieventris,

P. brevispinus, and P. placidus are the most studied and

most common.Almost all current research on Podisus spp. has focused on P. maculiventris, particularly its role in

soybean agroecosystems (see Irwin and Shepard 1980; Biever

et al. 1982). More studies examining situations in which a species will be an effective predator, biotypes, and

assessing the possibility of local specialization are needed. Important insights might arise from knowledge

of factors influencing oviposition, the range of a single

population, and cues used in prey location and capture.

One problem with using Podisus spp. in pest

suppression is their relative scarcity and tendency to

disperse rapidly when mass released.Kairomones, the

existence of which was indicated in McLain's (1979) study

of trail following behavior, might be synthesized to

retain predators in areas where they are needed.

Synthetic sex pheromones are a step towards attracting

predators to pest-infested fields.

Much work has focused on food consumption and

functional response. Emphasis should be given to

ascertaining what cues these insects use in searching for

prey. Research could focus on genetic improvement, for

example breeding a strain of P. maculiventris which 161

aggressively searches for and attacks prey, or one which has reduced flight abilities.

The Neotropical species of Podisus need to be revised.

From my examination of extralimital species, I believe that Supputius Distant will also need revisionary work.

Biological data are needed as well in order to assess the potential roles of Podisus spp. in South American pest management programs. TABLE 1. Summary of life history data forAptateticus and Podisus expressedin days. OW means overwintered and locationis where rearing was done. Standard abbrevi- ations are used for U. S. states. For Canadian provinces,BC means British Columbia, QE refers to Quebec,and NS is Nova Scotia. MX isMexico City.

Species Gen/ Egg I II III IV V Loca- Reference yr tion

A. brac- OW 9-15 14 13 17 19 BC Downes 1920 teatus

A. cyni- 1 OW 5 5 7 14 21 OH Whitmarsh 1916 cus

OW S16 46 37.g11 E7 S7 4312$11 - -- IL Stone 1939

1 OW 5.08 5.27 5.33 5.0 9.5 WI Jones and Coppel 1963

A. line------5-7 4-6 10-13 8-12 MX Plummer and Landis olatus 1932

P. acu- 5.1 2.6 3.8 2.9 3.4 5.1 AZ Stoner et al. 1974b tissimus

P. brevi- 1 15.35 7.75 7.71 7.83 11.94 23.82 QE Tostowaryk 1971 spinus

-6.5 3.9 4.1 3.9 5.0 7.6 WI Coppel and Jones 1962 TABLE 1. Continued

Species Gen/Egg I II III IV V Loca-Reference yr tion

b P. macu- 2.8- 3.0- 3.5- 6.0- OH Landis 1937 .... liventris 5.9 5.3 5.8 9.0

3 4.6 3.4 4.7 4.0 4.4 6.1 IL Esselbaugh 1948

1? 9.6 4.5 5.0 4.0 4.4 7.5 WI Coppel and Jones 1962

5.3c 4.9 5.4 4.8 4.4 8.9 QE Mukessji and LeRoux 1965

7.0 5.5 4.8 4.6 5.6 8.2 AR Warren and Wallis 1971

5.0 4.0 5.0 3.32 3.68 5.89 FLe Richman and Whitcomb 1978

5.0 4.0 4.85 3.15 3.58 6.11 FL Richman and Whitcomb 1978

6.5 4.0 5.0 4.0 4.0 7.5 OR present study TABLE 1. Continued

Species Gen/ Egg I II III IV V Loca-Reference yr tion

+ + + + P. pla- 2 -10 -7 -12 -10 - -- 21 MA Kirkland 1898 cidus

1 6.8 6.1 6.2 4.3 5.7 9.4 WI Coppel and Jones 1962

? 6.0 5.4 4.6 4.9 4.6 7.6 MO Oetting and Yonke 1971

P. saq- 12 11 9-13 8-14 7-15 13-17 MX Plummer and Landis itta 1932

P. serie- 3 8.0 ±4 8-10 5-6 ------MA Kirkland 1896 ventris

1 13.2 7.7 7.9 7.2 8.7 15.2 NS Prebble 1933

1 6.3 5.5 4.7 4.2 4.7 9.0 WI Coppel and Jones 1962

d aData are for 26.1°C. Means of data reported. b e Ranges given reflect development time on different hosts. Laboratory. c f Mean for 10 egg masses. Environator at 27°C. TABLE 2. Prey records of Apateticus and Podisus.

Species Prey Common Name Reference

A. bracteatus ORTHOPTERA Phasmatidae Diapheromera femorata (Say) Walkingstick Kirkland 1898

COLEOPTERA Chrysomelidae Calliqrapha elegans (Olivier) Beaulne 1939

Trirhabda borealis Blake Evans and Root 1980

Trirhabda yin:rata LeConte Ibid.

LEPIDOPTERA Arctiidae Hyphantria cunea (Drury) Fall webworm Morris 1972

Geometridae Lambdina fiscellaria sominaria Western oak (Huist) looper Downes 1920

Lasiocampidae Malacosoma sp. Tent caterpillars Ibid. TABLE 2, Continued

Species Prey Common Name Reference

A. bracteatus Noctuidae Spodoptera eridania (Cramer) Southern armyworm Evans and Root 1980

Pieridae Neophasia menapia (Felder and Pine butterfly Kirkland Felder) 1898

HYMENOPTERA Tenthredinidae Pristiphora erichsonii (Hartig) Larch sawfly Ives 1967, Muidrew 1955

Trichiocampus viminalis (Fallen) Downes 1920

A. cynicus ORTHOPTERA Phasmatidae Diapheromera femorata (Say) Walkingstick Kirkland 1898

HETEROPTERA Pentatomidae Podisus serieventris Uhler Kirkland 1898

COLEOPTERA Chrysomelidae Leptinotarsa decemlineata (Say) Colorado potato Kirkland beetle 1898

rn TABLE 2. Continued

Species Prey Common Name Reference

A. cynicus LEPIDOPTERA Arctiidae Halisodota caryae (Harris) Hickory tussock Whitmarsh moth 1916 Lasiocampidae Malacosoma americanum (Fabricius) Eastern tent Stone 1939 caterpillar Lymantriidae Lymantria dispar (L.) Gypsy moth Ibid. Noctuidae Alabama agrillacea (Hubner) Cotton leafworm Kirkland 1898 Datana anqusii Grote & Robinson Whitmarsh 1916

D. integerrima Grote & Robinson Walnut caterpillar Ibid.

D. ministra (Drury) Yellowheaded caterpillar Ibid. TABLE 2. Continued

Species Prey Common Name Reference

A. cynicus Saturniidae Callosamia promethea (Drury) Promethea moth Whitmarsh 1916

Hyalophora cecropia (L.) Cecropia moth Blatchley 1926 achemon (Drury) Achemon sphinx Whitmarsh 1916 HYMENOPTERA Diprionidae Diprion similis (Hartig) Introduced pine Jones and sawfly Coppel 1963 Tenthredinidae Nematus ribesii (Scopoli) Imported Kirkland currantworm 1898

Pristiphora erichsonii (Hartig) Larch sawfly Muldrew 1955 A. lineolatus HETEROPTERA Pentatomidae Apateticus lineolatus (Herrich - Plummer and Schaeffer) Landis 1932 COLEOPTERA Coccinellidae Epilachna varivestis Mulsant Mexican bean Ibid. (3) beetle co TABLE 2, Continued

Species Prey Common Name Reference

P. acutissimus LEPIDOPTERA Noctuidae Spodoptera exiqua (Htibner) Beet armyworm Stoner et al. 1974

P. brevispinus LEPIDOPTERA Geometridae (See McPherson Alsophila pometaria (Harris) Fall cankerworm Sherman 1982) 1921 Notodontidae Heterocampa quttivitta (Walker) Saddled prominent Hitchings 1910, Dudley 1919, Fisher 1970

Heterocampa spp. Fisher 1970

Olethreutidae Epinotia aceriella (Clemens) Maple trumpet Cote and skeletonizer Allen 1973 Laspereysia ptosamul (L.) Codling moth LeRoux 1960

P. fretus HYMENOPTERA Diprionidae Neodiprion excitans Rohwer Blackheaded pine Hetrick sawfly 1959 TABLE 2. Continued

Species Prey Common Name Reference

P. maculiven- HETEROPTERA tris Coreidae Anasa tristis (DeGeer) Squash bug Beard 1936 (See McPherson 1982) HOMOPTERA Cicadidae Maqicicada septendecim (L.) Periodical cicada Balduf 1943 Chrysopa oculata Say Goldeneye lacewing Ibid. COLEOPTERA Chrysomelidae Aqasicles hyqrophila Selman & Vogt Maddox et al. 1971

Trirhabda adela Blake Balduf 1943

T. borealis Blake Evans 1982a

T. virqata LeConte Ibid.

Trirhabda sp. Balduf 1943 Coccinellidae Hippodamia sp. Ibid.

0--J TABLE 2. Continued

Species Prey Common Name Reference 1111. P. maculiven- LEPIDOPTERA tris Dioptidae Phryclanidia californica Packard California oakwormBurke and Herbert 1920, Harville 1955, Pinnock and Milstead 1971 Noctuidae Anomis erosa Hubner Creighton 1936 eqena (Guenee) Bean leafskeleton- Genung 1960 izer

Spodoptera eridania (Cramer) Southern armyworm Evans 1982a DIPTERA Empididae Empis clausa Coquillett Balduf 1943 HYMENOPTERA Diprionidae Gilpinia hercyniae (Hartig) European spruce MacAloney sawfly 1936, Plumb 1936

Neodiprion swainei Middleton Swaine jack pine Becker and 1- sawfly Benjamin 1964 TABLE 2. Continued

Species Prey Common Name Reference

P. maculiven- Braconidae tris pas sp. Balduf 1943 P. mucronatus COLEOPTERA Chrysomelidae Agasicles =mai..a Selman & Vogt Maddox et al. 1971 LEPIDOPTERA Ctenuchidae Lymire edwardsii (Grote) Genung 1959 Noctuidae Autoplusia eciena (Guenee) Bean leafskeleton-Genung 1960 izer

Caterpillars on crucifers Hayslip et al. 1953 P. placidus HOMOPTERA Aphididae Acyrthosiphon pisum (Harris) Pea Wheeler 1977 COLEOPTERA Chrysomelidae Baliosus ruber (Walker) Basswood leafminerWest and Lothian 1948, Pyrrhalta luteola (Muller) Elm leaf beetle Torre Bueno 1939 TABLE 2. Continued

Species Prey Common Name Reference

P. placidus Coccinellidae Coleomegilla maculata (DeGeer) Wheeler 1977

Curculionidae Hypera postica (Gyllenhal) Alfalfa Ibid.

LEPIDOPTERA Arctiidae Hyphantria cunea (Drury) Fall webworm Kirkland 1898, Sulli- van and Green 1950, TadiC 1963, Warren and Bjegovi6 1972

Geornetridae Paleacrita vernata (Peck) Spring cankerworm LeRoux 1960 Lasiocampidae Malacosoma disstria Hubner Forest tent Kirkland caterpillar 1898

Malacosoma spp. Tent caterpillars Ibid. Lymantriidae Lymantria dispar (L.) Gypsy moth Kirkland 1898

Orgyia definita (Packard) Ibid. TABLE 2. Continued

Species Prey Common Name Reference

P. placidus Lymantriidae, cont'd Orqyia leucostiqmaJ. E. Smith) Whitemarked Kirkland tussock moth 1898

Noctuidae (Hufnagel) Black cutworm Oetting and Yonke 1971

Heliothis zea (Boddie) Corn earworm Ibid.

Lithophane antennata (Walker) Green fruitworm LeRoux 1960

Nymphalidae Nymphalis antiopa (L.) Mourningcloak Kirkland butterfly 1898

Oecophoridae Psilocarsis quercicella Clemens Carroll et al. 1979

P. reflexella Clemens

Pyralidae Galleria mellonella (L.) Greater wax moth Coppel and Jones 1962

Saturniidae Anisota senatoria (J. E. Smith) Orangestriped Felt 1905 oakworm 4 TABLE 2, Continued

Species Prey Common Name Reference

P. placidus Tortricidae Archips argyrospilus (Walker) Fruittree LeRoux 1960 leaf roller

DIPTERA Muscidae Musca autumnalis DeGeer Face fly Oetting and Yonke 1971

HYMENOPTERA Diprionidae Diprion similis (Hartig) Introduced pine Coppel and sawfly Jones 1962

Neodiprion pinetum (Norton) White pine sawfly Rauf and Benjamin 1980

Tenthredinidae Nematus ribesii (Scopoli) Imported Saunders currantworm 1870, Kirk land 1898, Fyles 1907

Pristiphora geniculata (Hartig) Mountain ash Forbes and sawfly Daviault 1964 TABLE 2. Continued

Species Prey Common Name Reference

P. saqitta HETEROPTERA Pentatomidae Podisus saqitta (Fabricius) Plummer and Landis 1932

COLEOPTERA Coccinellidae Epilachna varivestis Mulsant Mexican bean beetle Ibid.

P. serieven- LEPIDOPTERA tris Lymantriidae Orqyia pseudotsuqata Douglas-fir Dahlsten et (See McPherson (McDunnough) tussock moth al. 1977 1982) Pyralidae Galleria mellonella (L.) Greater wax moth Coppel and Jones 1962

HYMENOPTERA Diprionidae Gilpinia hercyniae (Hartig) European spruce Balch 1936, sawfly Reeks 1938 TABLE 3. Parasitoids attacking species of Podisus.

Species Parasitoid Stage Attacked

P. brevispinus SCELIONIDAE EGG Telenomus podisi Ashmead3

P. maculiventris ENCYRTIDAE EGG

OencyrtusSp.8

EUPELHIDAE EGG

Anastatus sp.1

SCELIONIDAE EGG

Cryon n. sp.1

Hadronatus sp.6

Telenomus dimmocki Ashmead3

T. podisi Ashmead3

Telenomus sp. 21

Telenomus sp. 41

Trissolcus basalis (Wollaston)1

T. euschistii (Ashmead)8 TABLE 3. Continued

Species ParasiLoid Stage Attacked

P. maculiventris SCELIONIDAE EGG Trissolcus podisi Ashmead3 T. thyantae Ashmead3

Trissolcus sp. B_1

TACHINIDAE NYMPH - ADULT Cylindromyia fumipennis Bigot'

Hemyda aurataEobineau-Desvoidy1 2 Trichopoda lanipes (Fabricius)-

P. placidus SCELIONIDAE EGG Telenomus persimilis Ashmead7

sacatta SCELTONIDAE EGG

Trissolcus sp.4 TABLE 3. Continued

Species Parasitoid Stage Attacked

P. serieventris SCELIONIDAE EGO Trissolcus podisi Ashmead5

1Buschman and Whitcomb 1980. Krombein et al. 1970.

2Fattig 1949. 6Potts 1891.

Girault 1907. 7Tadi 1968, 8 Gonzalez H. 1978. Yeargan 1979. 180

MAP 1: Distribution of Apateticus lineolatus (H. - S.) 181 182

MAP 2: Distribution of Apateticus marqiniventris (Stal). 183 184

MAP 3: Distribution of Apateticus bracteatus (Fitch).

186

MAP 4: Distribution of Apateticus anatarius Van Duzee.

188

MAP 5: Distribution of Apateticus cynicus (Say).

MAP 6: Distribution of Podisus mucronatus Uhler. SCALE to 1.1 L S 00 CU/ OW) nit OM( It PS I ..CopybiyhtDC PROJECTION 1939 by tha UnivolsIty of ChicoNo 1 192

MAP 7: Distribution of Podisus acutissimus St&l. 193 194

MAP 8: Distribution of Podisus maculiventris (Say).

196

MAP 9: Distribution of Podisus serieventris Uhler. Nrn MAP 10: Distribution of Podisus concirex (Staff). SCALE r.RC Copyright PROJECTION400 rsilg by E. University of Chicego 00 400 4,0.1114 200

MAP 11: Distribution of Podisus brevispinus, new species. 0N,--I 202

MAP 12: Distribution of Podisus fretus Olsen.

204

MAP 13: Distribution of Podisus placidus Uhler. 0NItn MAP 14: Distribution of Podisus saqitta (Fabricius). Leo SCALE .oa o. Copyright 19]9 by th linivrtityPROJECTION coo oo nn 01.1 I 1.5 fib. ago 208

FIGURE 1. Apateticus lineolatus (H. - S.).

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.5 mm.

d. Right clasper, dorsal view. Scale 0.5 mm.

e. Spermatheca. Scale 0.5 mm.

f. Sternite VI of female showing fuscous ring. Scale 1 mm.

FIGURE 2. Apateticus marginiventris (St&l).

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.5 mm.

d. Right clasper, dorsal view. Scale 0.5 mm.

e. Spermatheca. Scale 0.5 mm.

f. Ventral spine. Scale 1 mm. 209

lineolatus

a

marginiventris 210

FIGURE 3. Apateticus bracteatus (Fitch).

a. Anterolateral pronotal margins and humeri showing subacute and acute types, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 1 mm.

c. Right clasper, lateral view. Scale 0.5 mm.

d. Spermatheca. Scale 0.5 mm.

e. Female basal plates showing exposed triangular triangulin, ventro-caudal view. Scale 1 mm.

FIGURE 4, Apateticus anatarius Van Duzee.

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 1 mm.

c. Right clasper, lateral view. Scale 0.5 mm.

d. Spermatheca. Scale 0.5 mm.

e. Female basal plates showing V-shaped ridge on triangulin. Scale 1 mm. 211

a bracteatus

a anatarius 212

FIGURE 5. Apateticus cynicus (Say).

a. Ventral spine. Scale 1 mm.

b. Male genital cup, caudal view. Scale 1 mm.

c. Right clasper, lateral view. Scale 0.5 mm.

d. Spermatheca. Scale 0.5 mm.

e. Female basal plates showing square triangulin. Scale 1 mm.

FIGURE 6. Podisus mucronatus Uhler.

a. Anterolateral pronotal margins and humeral spines, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Proctiger showing C-shaped ridges. Scale 0.5 mm.

d. Right clasper, lateral view. Scale 0.25 mm.

e. Right clasper, dorsal view. Scale 0.25 mm.

f. Spermatheca. Scale 0.5 mm.

g. Female first gonocoxae, ventro-caudal view. Scale 0.5 mm. 213

mucronatus 214

FIGURE 7. Podisus acutissimus St&l.

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Male genital cup, dorsal view. Scale 0.5 mm.

d. Right clasper, mesal view. Scale 0.25 mm.

e. Right clasper, anterior view. Scale 0.25 mm.

f. Spermatheca. Scale 0.5 mm.

FIGURE 8. Podisus maculiventris (Say).

a. Anterolateral pronotal margins showing acute and spinose humeri. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.25 mm.

d. Right clasper, dorsal view. Scale 0.25 mm.

e. Spermatheca. Scale 0.5 mm.

f. Ventral spine. Scale 1 mm.

216

FIGURE 9. Podisus serieventris Uhler.

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.25 mm.

d. Spermatheca. Scale 0.5 mm.

e. Ventral spine. Scale 1 mm.

FIGURE 10. Podisus conclrex

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.25 mm.

d. Right clasper, dorsal view. Scale 0.25 mm.

e. Spermatheca. Scale 0.5 mm.

218

FIGURE 11. Podisus brevispinus, new species.

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.25 mm.

d. Right clasper, dorsal view. Scale 0.25 mm.

e. Spermatheca. Scale 0.5 mm.

f. Ventral spine. Scale 1 mm.

FIGURE 12. Podisus fretus Olsen.

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.25 mm.

d. Right clasper, dorsal view. Scale 0.25 mm.

e. Spermatheca. Scale 0.5 mm. je,6 I 220

FIGURE 13. Podisus placidus Uhler.

a. Anterolateral pronotal margins and humeri, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.25 mm.

d. Right clasper, dorsal view. Scale 0.25 mm.

e. Spermatheca. Scale 0.5 mm.

FIGURE 14. Podisus sagitta (Fabricius).

a. Anterolateral pronotal margins and humeral spines, dorsal view. Scale 1 mm.

b. Male genital cup, caudal view. Scale 0.5 mm.

c. Right clasper, lateral view. Scale 0.25 mm.

d. Right clasper, dorsal view. Scale 0.25 mm.

e. Spermatheca. Scale 0.5 mm. 221 222

FIGURE 15.

a. Rostrum, Supputius cincticeps (Staff), showing segment II equal to III + IV. Scale 1 mm.

b. Rostrum, P. maculiventris, showing segment II shorter than III + IV. Scale 1 mm.

c. Head, A. lineolatus, showing convergent juga typical of the genus. Dorsal view. Scale 1 mm.

d. Head, P. maculiventris, showing straight juga typical of the genus. Dorsal view. Scale 1 mm.

e. Conjunctival appendages, A. bracteatus, showing dorsal scierotized processes typical of the subgenus Apoecilus. Right view. Scale 0.5 mm.

f. Conjunctival appendages, A. lineolatus, showing scierotized tips typical of the subgenus Apateticus. Right view. Scale 0.5 mm.

g. Median penial lobes, A. bracteatus, left view. Scale 0.5 mm.

h. Median penial lobes, P. maculiventris, left view.

i. Right male genital plate, P. acutissimus. Scale 1 mm.

. Right male genital plate, P. maculiventris. Scale 0.25 mm.

k. Duct of metathoracic scent gland, P. acutissimus. Scale 1 mm.

1. Duct of metathoracic scent gland, P. maculiventris. Scale 1 mm. 223 15 224

FIGURE 16. Scanning electron micrograph of abdomen of Apateticus bracteatus, male, showing setose, rugulose areas on sternites IV - VI. 40 X. Photograph taken by Jerry Cassis. to N N 226

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* indicates not seen by the author. 244

APPENDICES 245

APPENDIX A

CHARACTERS USED IN THE CLADISTIC ANALYSES 246

DATA SETS FOR APATETICUS

* indicates a homoplasious character.

Data Set A-1: External

1. Juga: straight, equal to tylus (fig. 15d), (0); convergent, exceeding tylus (fig. 15c), (1).

2. Anterolateral margins of pronotum: concave, dentate (figs. la, 3a, 4a), (0); straight, edentate (fig. 2a), (1).

*3. Humeri: acute-spinose (fig. 8a), (0); blunt-subacute (figs. la, 2a, 3a, 4a), (1).

. Posterior pronotal angles of pronotum: blunt (figs. la, 2a), (0); acute-toothed (figs. 3a, 4a),(1).

*5. Ventral surface of corium, embolium: red (0); concolorous (1).

6. Membrane of hemelytra: with dark band (0); with dark spots (1); plain (2).

7. Body length: less than 12.5 mm (0); over 12.5 mm (1).

8. Setose, rugulose areas on abdominal venter of males: Absent (0); present (fig. 16), (1).

9. Abdominal venter: maculated (0); immaculate (1).

10. Ring-shaped black spot on sternite VI of females: absent (0); present (fig. if), (1).

11. Ventral spine: extending between metacoxae (fig. 5a, 8f), (0); very short (fig. 2f), (1).

12. Pleura: with 1 or 2 black spots (0); immaculate (1).

13. Duct of metathoracic scent gland: with black tip (0); plain (1).

*14. Tibiae: plain (0); banded apically and basally with black (1); with apical 2/3 black (2). 247

Data Set A-2: Male Genitalia

1. Genital cup: type B (fig. 8b), (0); type A (figs. 3b, 4b, 5b), (1).

*2. Genital cup: Type B; lateral lobes deflexed apically (fig. 8b), (0); type B; deeply concave between ventral margin and ventral surface (fig. lb), (1); type B; slightly concave between ventral margin and ventral surface (fig. 2b), (2); not type B (3).

3. Width of genital cup: 0.90 - 1.60 mm (0); 1.90 - 2.17 mm (1); 2.20 - 3.0+ mm (2).

. Proctiger without dorsomedian tubercle (figs. ib, 2b, 5b, 8b), (0); with dorsomedian tubercle (fig. 3b, 4b), (2).

5. Proctiger: not hourglass-shaped (fig. 8b), (0); hourglass-shaped (figs. lb, 2b), (1).

6. Claspers: type C, triangular-concave (fig. 8c), (0); type A, bifurcate (figs. 3c, 4c, 5c), (1); type B; bladelike, pointed (figs. lc, 2c), (2).

7. Claspers: not type A (0); type A with dorsal arm 3/4 length of ventral arm (fig. 3c), (1); type A with dorsal arm turned abruptly laterad, bluntly pointed (fig. 4c), (2); type A with dorsal arm much thinner than ventral arm (fig. 5c), (3).

8. Genital plates: quadrate with rounded corners (fig. 15j), (0); in shape of inverted L (figs. 3b, 4b, 5b), (1); in shape of inverted L with short base, wide stem (fig. ib, 2b), (2).

9. Conjunctival appendages; membranous (0); with sclerotized tips (fig. 15f), (1); with sclerotized dorsal process (fig. 15e), (2).

10. Median penial lobes: disk-shaped (fig. 15h), (0); long, tapered to blunt apex (fig. 15g), (1).

11. Thecal shield: of subequal size with theca (0); much larger than theca (1). 248

Data Set 3: Female Genitalia

1. Triangulin: not exposed (fig. 6g), (0); exposed (fig. 3e, 4e, 5e), (1).

2. Triangulin: not exposed (0); triangular (fig. 3e), (1); with V-shaped ridge (fig. 4e), (2); quadrate (fig. 5e), (3).

3. First gonocoxae: quadrate (fig. 4e, 5e), (0); triangular (fig. 3e), (1).

4. Spermathecal duct between dilation and pumping region: narrow (fig. 8e), (0); wide (fig. 4d),(1).

*5. Spermathecal duct: straight (figs. 2e, 4d), (0); looped, coiled (fig. le, 3d, 5d), (1).

6. Spermathecal bulb: elongate, curved (fig. 8e), (0); ovate (fig. 2c), (1); small, parallel-sided (fig.4d), (2).

7. Length of pumping region/length of sclerotized rod: 0.40 - 0.49 (0); 0.30 - 0.39 (1); 0.20 - 0.29(2).

Compiled Data Set for Final Cladogram

Character number in this set is followed by the number of that character from Data Set A-1, A-2, or A-3.

1. A-1-1 11. A-2-1

2. A-1-2 12. A-2-3

3. A-1-6 *13. A-2-6

4. A-1-7 14. A-2-7

5. A-1-8 *15 A-2-8

6. A-1-9 16. A-2-9

7. A-1-10 17. A-2-10

8. A-1-11 18. A-2-11

9. A-1-12 19. A-3-2

10. A-1-13 20. A-3-3 249

DATA SETS FOR PODISUS

* indicates a homoplasious character.

Data Set P-1: External

1. Length of rostral segment 2: exceeding 3 + 4 (fig. 15a), (0); less than 3 + 4 (fig. 15b), (1).

2. Rostral segment 4: concolorous with 1 - 3 (0); darker than 1 - 3 (1).

3. Length of pronotum anterior to humeri/total pronotal length: 0.57 - 0.62 (0); less than 0.57 (1).

*4. Abdominal venter maculated (0); immaculate (1).

5. Anterolateral margins of pronotum: concave (eg. fig. 7a, 8a), (0); straight (fig. 13a), (1).

6. Humeri: acute - spinose (fig. 8a), (0); blunt - subacute (figs. 9a, lla, 12a), (1).

7. Humeral spines: absent (fig. 13a), (0); entire(figs. 6a, 7a, 8a), (1); emarginate (fig. 14a), (2).

8. Transverse pronotal band: absent (0); present (1).

9. Scutellum: of one color (0); with apical half rufous - fuscous (1); with distinctivory tip (2).

10. Base of scutellum: plain or with irregular white areas (0); with 3 distinct ivory orrufous glabrous spots (1).

11. Ventral surface of corium, embolium: red (0); concolorous (1).

*12. Membrane of hemelytra: with dark band (0); plain (1).

13. Duct of metathoracic scent gland: curved (fig. 151), (0); straight (fig. 15k), (1). 250

*14. Ventral spine: extending between metacoxae (fig. 8f) (0); short, extending at most to posterior border of metacoxae (figs. 9e, 11f), (1); extending toanterior border of metacoxae (fig. 5a), (2).

15. Abdominal venter with fine (0) or coarse (1) punctation.

Data Set P-2: Male Genitalia

1. Genital cup: type A (figs. 3b, 4b, 5b),(0); type B (figs. 8b, 9b, 10b, 11b, 12b, 13b, 14b),(1); type C (fig. 7b), (2).

*2. Genital cup: not type B (0); type B withapically rounded lateral lobes (fig. 10b), (1); typeB with apically deflexed lateral lobes (figs. 8b, 9b,lib, 12b, 14b), (2).

3. Width of genital capsule: 0.90 - 1.60 mm (0); 1.65 - 1.85 mm (1).

*4. Area between ventral border and ventral surfaceof genital cup: flattened to slightly concave(figs, 6b, 13b), (0); with median depression belowlip-like ventral border (figs. 8b, 9b, 11b, 12b,14b), (1); narrow, concave (fig.10b), (2); otherwise (fig.7b), (3).

5. Proctiger: plain (eg. figs. 8b, 9b), (0); with C- shaped ridges dorsally, curvingmedially in basal quarter (fig. 6c), (1).

6. Claspers: type D (fig. 6d), (0); type E(fig. 7d), (1); type C (figs. 8c, 9c, 10c, 11c, 12c,13c, 14c), (2).

7. Claspers: not type C (0); type Cwith apically tapered, blunt dorsal and lateral angles(figs. 8c, 9c), (1); type C with basal half blunt, deep;slightly concave laterally (fig.10c), (2); type C with blunt dorsal and lateral apices and roundedlateral concavity (fig. 11c), (3); type C, widedorsally with narrow, laterally tapered ventralportion (fig. 12c), (4); type C, very narrow dorsally, withsubapical tooth on wider ventral portion(fig. 13c), (5); type C, narrow dorsally with wider,bluntly rounded ventral part (fig. 14c),(6). 251

8. Claspers type D with lateral arm pointed apically (0), with a V-shaped notch (fig. 6d), (1), or otherwise (2).

9. Genital plates: quadrate with rounded corners, lateral indentation, and convex mesally (fig.15j), (0); quadrate with mesal margin cut away, tuberculate (fig. 15i), (1).

Data Set P-3: Female Genitalia

*1. First gonocoxae: smooth (0); rugose (fig. 6g), (1).

*2. Posterior margin of first gonocoxae: undulant (fig. 6g), (0); straight (1).

*3. Spermathecal bulb: round (fig. 13e), (0); ovate (fig. 7f), (1); elongate - curved (figs. 8e, 9d, 12e), (2); spindle-shaped (fig. 14e), (3); truncate - elliptical (fig. lie), (4); elliptical (fig. 6f); (5).

4. Length of pumping region/length of sclerotized rod: 0.20 - 0.29 (0); 0.30 - 0.39 (1); 0.40 - 0.49(2); 0.50 - 0.59 (3).

*5. Proximal flange: small (figs. 6f, 12e, 13e), (0); prominent (figs. 8e, 9d), (1).

*6. Distal flange: small (figs. 13e, 14e), (0); prominent (figs. 7f, 8e, 9d), (1).

Compiled Data Set for Final Cladogram

Character number in this set is followed by the number of that character from Data Set P-1, P-2, or P-3.

1. P-1-1 7. P-1-10 12. P-2-5

2. P-1-2 8. P-1-11 13. P-2-6

3. P-1-3 9. P-1-15 14. P-2-7

4. P-1-5 10. P-2-1 15. P-2-8

5. P-1-8 11. P-2-3 16. P-2-9

*6. P-1-9 252

APPENDIX B

CLADOGRAMS SHOWING THE RELATIONSHIPS AMONG

THE SPECIES OF APATETICUS AND PODISUS 253

Coding of the c--coci,=s in th,,, rladogram

1. Genus Apatetcus

MACU: Poclisus naculiventris (outgroup)

LINE: lineolatus

MA?G: nar7nivni-ris

nr,,AC'e bracteatus anatarius

CYNI: cvnicus

2. Genus Podisus

SUCI: Sumnutius cincticeps (outcrouo

rucronatus,

ACUT: acutisirlzs

MACU: rarul'ven4-r4s

serieventris

GONG: concrex

hrovis-'n'1.7

FRET: fretus

-DT r3lacidus

SAGI: sacitta 254

Cladogram 1. Species of Apateticus based on external characters (Data Set A-1). Bold numbers indicate homoplasious characters; light numbers are nodes.

Cladogram 2. Species of Apateticus based on characters of the male genitalia (Data Set A-2). Bold numbers indicate homoplasious characters; light numbers are nodes.

Cladogram 3. Species of Apateticus based on characters of the female genitalia (Data SetA-3). Bold numbers indicate homoplasious characters; light numbers are nodes. 255

BRAC 14 CYNI 40 - --3 5o 2 t ANAT 3 4t14-2- MARS 4 34. 51 141 LINE 5 NACU

ANAT

1 CYNI40 23 41 BRAG 4 NARG 4 24 3 4 LINE 5 NACU

CYNI

ANAT 50

BRAG

LINE 54 4

MARG 5

NACU Cladogram 4. Species of Apateticus based on the compiled data set(Data Sets A-1, A-2, and A-3). Numbers indicate correlated characters. LGZ

maculiventris isus Pod 258

Cladogram 5. Species of Podisus based on external characters (Data Set P-1). Bold numbers indicate homoplasious characters; light numbers are nodes.

Cladogram 6. Species of Podisus based on characters of the male genitalia (Data Set P-2). Bold numbers indicate homoplasious characters; light numbers are nodes. 259

MUCR B ACUT 40 142

CMG 3 NACU 121 3 SAGI 6 FLAG121 6 6 FRET 6 6 SERI 6 1141 2

BREV 9 SUCI

SAGI 4 FLAG 21 40 3 FRET

REV

cono2i I 2 6 SERI 6

NACU

ACUT 43 7 22

MUCR 9

SUCI 260

Cladogram 7. Species of Podisus based on characters of the female genitalia (Data SetP-3). Bold numbers indicate homoplasious characters; light numbers are nodes. 261

SAGI 60 2 33 7 BRED 34 4 mACU 4 5,6, ACUT 31 4 SERI 5 MUCR 120-35 32

FRET 6 PLAC 2 6 GONG 1151 ai 9 SUCI Cladogram 8. Species of Podisus based on thecompiled data set (Data Sets P-1, P-2, and P-3). Numbers indicate correlated characters. £9Z

^Co

cincticeps Supputius