Great Lakes Entomologist the Grea T Lakes E N Omo L O G Is T Published by the Michigan Entomological Society Vol

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The Great Lakes Entomologist The Grea t Lakes E n omo l o g is t Published by the Michigan Entomological Society Vol. 45, Nos. 3 & 4 Fall/Winter 2012

Volume 45 Nos. 3 & 4 ISSN 0090-0222

Table of Contents The Scholar, Teacher, and Mentor: A Tribute to Dr. J. E. McPherson...... i Great Lakes Dr. J. E. McPherson, Educator and Researcher Extraordinaire: Biographical Sketch and List of Publications Thomas J. Henry...... 111 J.E. McPherson – A Career of Exemplary Service and Contributions to the Entomological Entomologist Society of America George G. Kennedy...... 124 Mcphersonarcys, a New Genus for Pentatoma aequalis Say (Heteroptera: Pentatomidae) Donald B. Thomas...... 127 The Stink Bugs (Hemiptera: Heteroptera: Pentatomidae) of Missouri Robert W. Sites, Kristin B. Simpson, and Diane L. Wood...... 134 Tymbal Morphology and Co-occurrence of Spartina Sap-feeding Insects (Hemiptera: Auchenorrhyncha) Stephen W. Wilson...... 164 Pentatomoidea (Hemiptera: Pentatomidae, Scutelleridae) Associated with the Dioecious Shrub Florida Rosemary, Ceratiola ericoides (Ericaceae) A. G. Wheeler, Jr...... 183 An Annotated Checklist of the Stink Bugs (Heteroptera: Pentatomidae) of New Mexico Vol. 45, Nos. 3 & 4 C. Scott Bundy...... 196 The Pentatomidae, or Stink Bugs, of Kansas with a Key to Species (Hemiptera: Heteroptera) Richard J. Packauskas...... 210 Annotated Checklist of the Pentatomidae (Heteroptera) of Connecticut Jane E. O’Donnell, and Carl W. Schaefer...... 220 The Genus Sphyrocoris Mayr (Heteroptera: Scutelleridae: Pachycorinae) J. E. Eger, Jr...... 235 The Stink Bugs (Hemiptera: Heteroptera: Pentatomidae) of Washington State Richard S. Zack, Peter J. Landolt, and Joseph E. Munyaneza...... 251 An Updated Synopsis of the Pentatomoidea (Heteroptera) of Michigan D. R. Swanson...... 263 The Heteroptera (Hemiptera) of North Dakota I: Pentatomomorpha: Pentatomoidea David A. Rider...... 312

Cover photo Published by Adult Mcphersonarcys aequalis (Say) Photo by Robert W. Sites The Michigan Entomological 2012 Society 2012THE GREATTHE LAKESGREAT LAKES ENTOMOLOGIST ENTOMOLOGIST 109 Published by the Michigan Entomological Society

Volume 45 Fall/Winter 2012 Nos. 3 & 4

SCHOLAR, TEACHER, AND MENTOR: A TRIBUTE TO DR. J. E. MCPHERSON

Dr. J. E. McPherson

It is with great pleasure that we dedicate this Festschrift to J. E. (Jay) McPherson on the occasion of his retirement. Jay has been one of the most prolific authors and a regular and insightful reviewer for this journal for several decades. He has also provided much-appreciated editorial expertise and advice over the years. We thank the authors who worked diligently during the past year to bring together this series of papers on Heteroptera, primarily Pentatomidae, the focus of Jay’s research to which he has contributed immense knowledge over the course of his career. This celebratory issue is a testament to the influence that Jay has had over his colleagues and former students and the esteem they feel for him. Congratulations and best wishes, Jay!

Governing Board of the Michigan Entomologocial Society

(photo used with permission from the Entomological Society of America) (i) 110 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

(ii) 2012 THE GREAT LAKES ENTOMOLOGIST 111 Dr. J. E. McPherson, Educator and Researcher Extraordinaire: Biographical Sketch and List of Publications Thomas J. Henry1 Like many outstanding naturalists, John E. (“Jay”) McPherson grew up with a strong interest in the natural world, especially insects. This innate curiosity led him to enroll as a zoology major at San Diego State University in 1959. Upon completion of his undergraduate degree, he continued on to pursue his interest in insect biology, completing a Master’s thesis on the life history and morphology of a poorly known species of Notonectidae. Shortly thereafter, a teaching assistantship enabled him to pursue a Ph.D. at Michigan State Uni- versity in East Lansing, where his research involved distinguishing two closely related species of bark beetles. During this period, Jay also worked on various pest species, including the cereal leaf beetle, pine tip beetle, and pine cone beetle. In 1969, Jay took a position at Southern Illinois University (SIU), Car- bondale (SIU) as an assistant professor. There he developed a broad and successful educational program, in which he taught numerous courses, including general zoology, introductory and advanced entomology, aquatic entomology, and pest control. He soon was promoted to Associate Professor (1974) and Full Professor (1979). Over his career, Jay (Fig. 1) has taught thousands of students, directed three Ph.D. and 23 Master’s students, and served on more than 40 thesis

Figure 1. J. E. McPherson in his “McPherson Lab Survivor” T-shirt at SIU, June 2008 (photo courtesy of Robert W. Sites).

1Systematic Entomology Laboratory, Agricultural Research Service, United States Department of Agriculture, c/o National Museum of Natural History, MRC-168, Smithsonian Institution, Washington, D. C. 20013-7013 (e-mail: thomas.henry@ars. usda.gov). 112 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 committees. His students have pursued a wide range of research topics; as a heteropterist, I have followed most closely those who studied the Hemiptera. Those who have gone on to have outstanding careers of their own, include Jeff (Jeffrey D.) Bradshaw [reduviid biology and morphology], Scott (C. Scott) Bundy [pentatomoid biology and morphology], Steve (Steven L.) Keffer [Nepidae], Rich (Richard J.) Packauskas [Coreidae], Bob (Robert W.) Sites [Naucoridae], Steve (Stephen J.) Taylor [biology and taxonomy of Gerromorpha and Aradidae), and Steve (Stephen W.) Wilson [Fulgoroidea]. As testament to his success and dedication to teaching, he received in 1993 the North Central Branch, ESA Distinguished Achievement Award in Teaching and, in 1996, earned the Out- standing Teacher Award from the College of Science at SIU. In addition to Jay’s heavy involvement in teaching, he has been extraor- dinarily active in his university and scientific societies. He has served on more than 46 committees at SIU. He has been chairman of the Graduate and Undergraduate Student committees, served on numerous search committees for departmental staff positions, been acting Chairman of the Department, and was long the Assistant Chair for the Department of Zoology, beginning in 1980 until his retirement. At the college and university levels, Jay served on more than 20 committees, including the Biological Sciences Advisory Committee, College of Science Dean Search Committee (Chair), Blue Ribbon Committee to consider the future of SIU, and Outstanding Dissertation Award Committee for the University. Jay’s contributions to scientific societies are many, particularly the Ento- mological Society of America (Fig. 2), where he helped create the ever-popular Linnaean Games, served on the Governing Board for six years (two terms), served as editor of the American Entomologist for nine years (see Wheeler 2002. American Entomologist: Summer, p. 66), and was vice president (2001) and president of the Society (2002). That aspect of his career is covered in an article by George Kennedy in this issue honoring Dr. McPherson. Despite his focus on teaching and university affairs and his remarkable societal accomplishments, Jay has somehow managed to maintain an impressively productive research program. During his career, he has produced more than 180 scientific publications, primarily on the biology, life history, and immature stages of true bugs, especially the Pentatomoidea, Reduviidae, and several families of aquatic bugs, including Belostomatidae, Gerridae, Nepidae, and Notonectidae. He has published detailed descriptions of 193 nymphal stages and 39 eggs of Heteroptera in 21 families that serve as models for others to follow. He also published a series of classic papers (e.g., McPherson 1975, 1976, 1979) documenting the effects of photoperiod on the adult morphology of Euschistus tristigmus (Say). More recently, he has focused on certain other Pentatomidae, including studies on the comparative genitalic morphology of Brochymena and Parabrochymena (McPherson 2007), the U.S. species of Euschistus (McPherson and Ahmad 2008), two closely related species of Mecidea (McPherson et al. 2009), and three species of Murgantia (McPherson and Ahmad 2008), the last of which resulted in the discovery of a new species (McPherson and Ahmad 2011). His recent interests also involve the Reduviidae, which include the description of the new harpacto- rine genus Parasinea to accommodate a new species from Colombia and a previously known species transferred from Sinea (McPherson and Ahmad 2011), and clarification of the identity of Sinea sericea Hussey (McPherson 2012). Among his most outstanding contributions are his book chapter (Panizzi et al. 2000) in Panizzi and Schaefer’s Heteroptera of Economic Importance on the economically important species of Pentatomoidea of the world, treating about 40 genera and 58 species, and two important books, the Pentatomoidea of Northeastern North America (McPherson 1982), with keys and an exhaustive literature review of 58 genera and 120 species and subspecies, and the Stink Bugs of Economic Impor- tance in America North of Mexico (McPherson and McPherson 2000), with keys and overviews of 23 genera and 45 species. 2012 THE GREAT LAKES ENTOMOLOGIST 113

Figure 2. J. E. McPherson at the Entomological Society of America Annual Meeting, Reno, Nevada, November 2011 (photo courtesy of C. Scott Bundy).

Jay McPherson has had a remarkable career as an award-winning teacher, as a leader and active participant in ESA, including president in 2002, and as a scholarly researcher. Although Jay has retired (15 June 2012), a recent conversation that I had with him suggested little will change. He likely will continue teaching on a contractual basis, he certainly will stay active in ESA, and clearly his research program shows little sign of slowing. Congratulations Dr. McPherson on your retirement and many accomplishments. We applaud your outstanding career and look forward to many more productive years.

List of Publications by J. E. McPherson 1. McPherson, J. E. 1965. Notes on the life history of Notonecta hoffmanni (Hemiptera: Notonectidae). Pan-Pacific Entomologist 41: 86–89. 2. McPherson, J. E. 1966. Notes on the laboratory rearing of Notonecta hoffmanni (He- miptera: Notonectidae). Pan-Pacific Entomologist 42: 54–56. 3. McPherson, J. E. 1967. Brief descriptions of the external anatomy of the various stages of Notonecta hoffmanni (Hemiptera: Notonectidae). Pan-Pacific Entomolo- gist 43: 117–121. 4. McPherson, J. E., and L. F. Wilson. 1967. Grasshoppers feeding on red pine trees in Michigan (Orthoptera: Acrididae). The Michigan Entomologist 1: 249–252. 5. McPherson, J. E., L. F. Wilson, and F. W. Stehr. 1970. A comparison between Conoph- thorus shoot-infesting beetles and Conophthorus resinosae (Coleoptera: Scolytidae). I, Comparative life history studies in Michigan. The Canadian Entomologist 102: 1008–1015. 114 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

6. McPherson, J. E., F. W. Stehr, and L. F. Wilson. 1970. A comparison between Conoph- thorus shoot-infesting beetles and Conophthorus resinosae (Coleoptera: Scolytidae). II, Reciprocal host and resin toxicity tests; with description of a new species. The Canadian Entomologist 102: 1016–1022. 7. McPherson, J. E. 1970. A key and annotated list of the Scutelleroidea of Michigan (Hemiptera). The Michigan Entomologist 3: 34–63. 8. McPherson, J. E. 1971. Notes on the laboratory rearing of Corimelaena lateralis lateralis (Hemiptera: Corimelaenidae) on wild carrot. Annals of the Entomological Society of America 64: 313–314. 9. McPherson J. E., and J. F. Walt. 1971. The first record in Illinois of a population of Stethaulax marmoratus (Say) (Hemiptera: Scutelleridae) with information on life history. Transactions of the Illinois State Academy of Science 64 :198–200. 10. McPherson, J. E. 1971. Laboratory rearing of Euschistus tristigmus tristigmus. Journal of Economic Entomology 64: 1339–1340. 11. McPherson, J. E. 1972. Life history of Corimelaena lateralis lateralis (Hemiptera: Thyreocoridae) with descriptions of immature stages and list of other species of Scutelleroidea found with it on wild carrot. Annals of the Entomological Society of America 65: 906–911. 12. McPherson, J. E. 1972. Laboratory rearing of Trichopepla semivittata (Hemiptera: Pen- tatomidae) on wild carrot. Annals of the Entomological Society of America 65: 985–987. 13. Walt, J. F., and J. E. McPherson. 1972. Laboratory rearing of Stethaulax marmoratus (Hemiptera: Scutelleridae). Annals of the Entomological Society of America 65: 1242–1243. 14. Walt, J. F., and J. E. McPherson. 1973. Descriptions of immature stages of Stethaulax marmorata (Hemiptera: Scutelleridae) with notes on its life history. Annals of the Entomological Society of America 66: 1103–1107. 15. McPherson, J. E. 1974. The first record in Illinois of Aelia americana (Hemiptera: Pentatomidae). Transactions of the Illinois State Academy of Science 67: 151. 16. McPherson, J. E. 1974. Photoperiod effects in a southern Illinois population of the Euschistus tristigmus complex (Hemiptera: Pentatomidae). Annals of the Entomo- logical Society of America 67: 943–952. 17. McPherson, J. E. 1974. Notes on the biology of Mormidea lugens and Euschistus politus (Hemiptera: Pentatomidae) in southern Illinois. Annals of the Entomological Society of America 67: 940–942. 18. McPherson, J. E. 1974. Three negro bug state records for Illinois (Hemiptera: Corimelaenidae). Transactions of the Illinois State Academy of Science 67: 361–363. 19. McPherson, J. E., and J. P. Cuda. 1974. The first record in Illinois ofNezara viridula (Hemiptera: Pentatomidae). Transactions of the Illinois State Academy of Science 67: 461–462. 20. Vangeison, K. W., and J. E. McPherson. 1975. Life history and laboratory rearing of Proxys punctulatus (Hemiptera: Pentatomidae) with descriptions of immature stages. Annals of the Entomological Society of America 68: 25–30. 21. McPherson, J. E., and R. H. Mohlenbrock. 1975. The Scutelleroidea of the LaRue- Pine Hills Ecological Area. Proceedings of the North Central Branch, Entomological Society of America 29: 157. (Abstract) 22. McPherson, J. E. 1975. Life history of Euschistus tristigmus tristigmus (Hemiptera: Pentatomidae) with information on adult seasonal dimorphism. Annals of the En- tomological Society of America 68: 333–334. 23. McPherson, J. E., and K. W. Vangeison. 1975. Effects of photoperiod on a population of Euschistus tristigmus (Say) (Hemiptera: Pentatomidae) from Gainesville, Florida. Annals of the Entomological Society of America 68: 205–206. 2012 THE GREAT LAKES ENTOMOLOGIST 115

24. McPherson, J. E., and J. P. Cuda. 1975. The first record in Illinois of Prionosoma podopioides (Hemiptera: Pentatomidae). Journal of the Kansas Entomological Society 48: 371–373. 25. McPherson, J. E. 1975. Effects of developmental photoperiod on adult morphology in Euschistus tristigmus tristigmus (Say) (Hemiptera: Pentatomidae). Annals of the Entomological Society of America 68: 1107–1110. 26. Youther, M. L., and J. E. McPherson. 1975. A study of fecundity, fertility, and hatch in Euschistus servus (Hemiptera: Pentatomidae) with notes on precopulatory and copulatory behavior. Transactions of the Illinois State Academy of Science 68: 321–338. 27. Nixon, P. L., J. E. McPherson, and J. P. Cuda. 1975. A list of the Scutelleroidea (Hemiptera) collected on immature black walnut trees in southern Illinois with some notes on biology. Transactions of the Illinois State Academy of Science 68: 409–413. 28. McPherson, J. E. 1975. A confirming report of the presence in southern Illinois of Brochymena cariosa (Hemiptera: Pentatomidae). Transactions of the Illinois State Academy of Science 68: 438–439. 29. McPherson, J. E., and R. H. Mohlenbrock. 1976. A list of the Scutelleroidea of the LaRue-Pine Hills Ecological Area with notes on biology. The Great Lakes Entomolo- gist 9: 125–169. 30. McPherson, J. E. 1976. Effects of photoperiod on a population of Euschistus tristigmus (Hemiptera: Pentatomidae) from Storrs, CT. Annals of the Entomological Society of America 69: 691–694. 31. Cuda, J. P., and J. E. McPherson. 1976. Life history and laboratory rearing of Bro- chymena quadripustulata with descriptions of immature stages and additional notes on Brochymena arborea (Hemiptera: Pentatomidae). Annals of the Entomological Society of America 69: 977–983. 32. McPherson, J. E. 1976. The first record in Illinois ofBrochymena punctata punctata (Hemiptera: Pentatomidae). Transactions of the Illinois State Academy of Science 69: 281–282. 33. McPherson, J. E. 1976. Notes on the biology of Cosmopepla bimaculata (Hemiptera: Pentatomidae) in southern Illinois. Transactions of the Illinois State Academy of Science 9: 362–366. 34. McPherson, J. E. 1977. Notes on the biology of Thyanta calceata (Hemiptera: Pen- tatomidae) with information on adult seasonal dimorphism. Annals of the Entomo- logical Society of America 70: 370–372. 35. McPherson, J. E. 1977. Effects of developmental photoperiod on adult color and pubescence in Thyanta calceata (Hemiptera: Pentatomidae) with information on ability of adults to change color. Annals of the Entomological Society of America 70: 373–376. 36. Nixon, P. L., and J. E. McPherson. 1977. An annotated list of phytophagous insects collected on immature black walnut trees in southern Illinois. The Great Lakes Entomologist 10: 211–222. 37. McPherson, J. E. 1977. The first report of the occurrence ofSymphylus caribbeanus (Hemiptera: Scutelleridae) in Texas. Journal of the Kansas Entomological Society 50: 529–530. 38. McPherson, J. E. 1977. A confirming report of the presence in southern Illinois of Banasa sordida (Hemiptera: Pentatomidae). Transactions of the Illinois State Academy of Science 70: 108–109. 39. McPherson, J. E. 1977. Three new scutelleroid state records for Illinois (Hemiptera). Transactions of the Illinois State Academy of Science 70: 388–390. 40. McPherson, J. E. 1978. Sensitivity of immature Thyanta calceata (Hemiptera: Pentatomidae) to photoperiod as reflected by adult color and pubescence. The Great Lakes Entomologist 11: 71–76. 116 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

41. McPherson, J. E. 1978. A reexamination of the Galgupha aterrima Malloch types (He- miptera: Thyreocoridae). Journal of the Kansas Entomological Society 51: 403–404. 42. McPherson, J. E., and R. I. Sailer. 1978. A new species of Corimelaena (Hemiptera: Thyreocoridae) from America north of Mexico. Journal of the Kansas Entomological Society Soc. 51: 516–520. 43. McPherson, J. E. 1978. Effects of various photoperiods on color and pubescence in Thyanta calceata (Hemiptera: Pentatomidae). The Great Lakes Entomologist 11: 155–158. 44. McPherson, J. E. 1978. A list of the Scutelleroidea (Hemiptera) of southern Illinois. The Great Lakes Entomologist 11: 159–162. 45. McPherson, J. E. 1979. Effects of various photoperiods on morphology in Euschistus tristigmus tristigmus (Hemiptera: Pentatomidae). The Great Lakes Entomologist 12: 23–26. 46. McPherson, J. E. 1979. Additions and corrections to the list of Michigan Pentato- moidea (Hemiptera). The Great Lakes Entomologist 12: 27–29. 47. McPherson, J. E. 1979. Effects of continuous and split developmental photophases during each 24 hour period on adult morphology in Euschistus tristigmus tristigmus (Hemiptera: Pentatomidae). The Great Lakes Entomologist 12: 79–81. 48. McPherson, J. E. 1979. Effects of various photoperiods on color and pubescence in Thyanta pallidovirens accerra (Hemiptera: Pentatomidae). The Great Lakes Entomologist 12: 83–84. 49. Wilson, S. W., and J. E. McPherson. 1979. A preliminary report on the bionomics of Fulgoroidea on black walnut with emphasis on Anormensis septentrionalis (Spinola) (Homoptera). USDA Forest Service General Technical Report NC-52: 29–34. 50. McPherson, J. E. 1979. A revised list of the Pentatomoidea of Illinois (Hemiptera). The Great Lakes Entomologist 12: 91–98. 51. Wilson, S. W., and J. E. McPherson. 1979. Notes on the occurrence of Epipyrops barberiana Dyar in southern Illinois (Lepidoptera: Epipyropidae). The Great Lakes Entomologist 12: 189–190. 52. Wilson, S. W., and J. E. McPherson. 1979. The first record ofMegamelus paleatus in Illinois (Homoptera: Fulgoroidea: Delphacidae). The Great Lakes Entomologist 12: 177. 53. Wilson, S. W., and J. E. McPherson. 1979. The first report of the occurrence ofOecleus chrisjohni and O. epetrion in Illinois (Homoptera: Fulgoroidea: Cixiidae). The Great Lakes Entomologist 12: 227–228. 54. Sites, R. W., and J. E. McPherson. 1979. The first record in southern Illinois of Polygonia progne (Lepidoptera: Nymphalidae). Transactions of the Illinois State Academy of Science 72: 93. 55. McPherson, J. E. 1980. The distribution of the Pentatomoidea in the northeastern quarter of the United States (Hemiptera). The Great Lakes Entomologist 13: 1–16. 56. McPherson, J. E. 1980. A list of the prey species of Podisus maculiventris (Hemiptera: Pentatomidae). The Great Lakes Entomologist 13: 17–24. 57. Wilson, S. W., and J. E. McPherson. 1980. A list of the Fulgoroidea (Homoptera) of southern Illinois. The Great Lakes Entomologist 13: 25–30. 58. Jones, W. A., and J. E. McPherson. 1980. The first report of the occurrence of acanthosomatids (Hemiptera: Acanthosomatidae) in South Carolina. Journal of the Georgia Entomological Society 15: 286–289. 59. Sites, R. W., and J. E. McPherson. 1980. A key to the butterflies of Illinois (Lepidop- tera: Papilionoidea). The Great Lakes Entomologist 13: 97–114. 60. McPherson, J. E. 1980. A peculiar specimen of Euschistus servus from North Carolina (Hemiptera: Pentatomidae). The Great Lakes Entomologist 13: 139–140. 2012 THE GREAT LAKES ENTOMOLOGIST 117

61. McPherson, J. E. 1980. The first report of the occurrence ofBrochymena carolinensis in Missouri (Hemiptera: Pentatomidae). The Great Lakes Entomologist 13: 168. 62. McPherson, J. E., and B. C. Weber. 1980. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 1. Pentatomoidea. The Great Lakes Entomologist 13: 177–183. 63. Wilson, S. W., and J. E. McPherson. 1980. Mixed species feeding assemblages of planthopper nymphs (Homoptera: Fulgoroidea). The Great Lakes Entomologist 13: 185. 64. Sites, R. W., and J. E. McPherson. 1981. The first record in southern Illinois of Celestrina ebenina (Lepidoptera: Lycaenidae). Transactions of the Illinois State Academy of Science 73(1)[1980]: 86. 65. Wilson, S. W., and J. E. McPherson. 1981. Keys to the planthoppers, or Fulgoroidea, of Illinois (Homoptera). Transactions of the Illinois State Academy of Science 73(2) [1980]: 1–61. 66. Clair, D. J., and J. E. McPherson. 1981. Effects of temperature on development and reproduction in Euschistus tristigmus tristigmus (Hemiptera: Pentatomidae) with notes on reproductive behavior. Transactions of the Illinois State Academy of Sci- ence 73(2)[1980]: 80–92. 67. Wilson, S. W., and J. E. McPherson. 1981. The distribution of the Fulgoroidea of the eastern United States (Homoptera). Transactions of the Illinois State Academy of Science 73(4)[1980]: 7–20. 68. Wilson, S. W., and J. E. McPherson. 1981. A list of the host plants of the Illinois Acanaloniidae and Flatidae (Homoptera: Fulgoroidea). Transactions of the Illinois State Academy of Science 73(4)[1980]: 21–29. 69. McPherson, J. E., and B. C. Weber. 1981. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 2. Coreoidea. The Great Lakes Ento- mologist 14: 11–13. 70. McPherson, J. E., and B. C. Weber. 1981. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 3. Reduvioidea. The Great Lakes Entomologist 14: 15–17. 71. McPherson, J. E., and B. C. Weber. 1981. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 4. Cimicoidea. The Great Lakes Ento- mologist 14: 19–22. 72. Wilson, S. W., and J. E. McPherson. 1981. Notes on the biology of Nersia florens (Homoptera: Fulgoroidea: Dictyopharidae) with descriptions of eggs, and first, second, and fifth instars. The Great Lakes Entomologist 14: 45–58. 73. Wilson, S. W., and J. E. McPherson. 1981. Ontogeny of the tibial spur in Megamelus davisi (Homoptera: Delphacidae) and its bearing on delphacid classification. The Great Lakes Entomologist 14: 49–50. 74. McPherson, J. E., and T. E. Vogt. 1981. The first report of the occurrence ofMecidea major (Hemiptera: Pentatomidae) in Illinois. The Great Lakes Entomologist 14: 70. 75. Sites, R. W., and J. E. McPherson. 1981. A list of the butterflies (Lepidoptera: Pa- pilionoidea) of the LaRue-Pine Hills Ecological Area. The Great Lakes Entomologist 14: 81–85. 76. Wilson, S. W., and J. E. McPherson. 1981. Life histories of Acanalonia bivittata and A.conica with descriptions of immature stages. Annals of the Entomological Society of America 74: 289–298. 77. Wilson, S. W., and J. E. McPherson. 1981. Life histories of Anormenis septentrionalis, Metcalfa pruinosa, and Ormenoides venusta with descriptions of immature stages. Annals of the Entomological Society of America 74: 299–311. 118 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

78. Wilson, S. W., and J. E. McPherson. 1981. Descriptions of the immature stages of Bruchomorpha oculata with notes on laboratory rearing. Annals of the Entomological Society of America 74: 341–344. 79. Wilson, S. W., and McPherson. 1981. Life history of Megamelus davisi with descriptions of immature stages. Annals of the Entomological Society of America 74: 345–350. 80. McPherson, J. E., and B. C. Weber. 1981. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 5. Lygaeoidea. The Great Lakes Ento- mologist 14: 133–136. 81. McPherson, J. E., and B. C. Weber. 1981. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 6. Tingidae and Aradidae. The Great Lakes Entomologist 14: 137–140. 82. Sites, R. W., and J. E. McPherson. 1981. The first records in Illinois of Heliconius charitonius (Lepidoptera: Heliconiidae) and Phoebis agarithe (Lepidoptera: Pieridae). The Great Lakes Entomologist 14: 205–206. 83. Sites, R. W., and J. E. McPherson. 1982. Life history and laboratory rearing of Se- hirus cinctus cinctus (Hemiptera: Cydnidae), with descriptions of immature stages. Annals of the Entomological Society of America 75: 210–215. 84. McPherson, J. E., and S. M. Paskewitz. 1982. Effects of continuous and split developmental photophases during each 24-hour period on adult color and pubescence in Thyanta calceata (Hemiptera: Pentatomidae). The Great Lakes Entomologist 15: 97–98. 85. Biehler, J. A., and J. E. McPherson. 1982. Life history and laboratory rearing of Galgupha ovalis (Hemiptera: Corimelaenidae), with descriptions of immature stages. Annals of the Entomological Society of America 75: 465–470. 86. McPherson, J. E., and S. M. Paskewitz. 1982. Effects of various split developmental photophases and constant light during each 24-hour period on adult morphology in Euschistus tristigmus tristigmus (Hemiptera: Pentatomidae). The Great Lakes Entomologist 15: 159–161. 87. Weber, B. C., and J. E. McPherson. 1982. The distribution of Xylosandrus germanus in America north of Mexico (Coleoptera: Scolytidae). The Great Lakes Entomologist 15: 171–174. 88. Weber, B. C., and J. E. McPherson. 1982. Annotated bibliography of the ambrosia beetle Xylosandrus germanus (Coleoptera: Scolytidae). The Great Lakes Entomolo- gist 15: 175–183. 89. Paskewitz S. M., and J. E. McPherson. 1982. Descriptions of nymphal instars of Thy- anta calceata (Hemiptera: Pentatomidae). The Great Lakes Entomologist 15: 231–235. 90. McPherson, J. E. 1982. The Pentatomoidea (Hemiptera) of northeastern North America with emphasis on the fauna of Illinois. Southern Illinois University Press, Carbondale and Edwardsville. 240 pp. 91. Weber, B. C., and J. E. McPherson. 1983. Life history of the ambrosia beetle Xylo- sandrus germanus (Coleoptera: Scolytidae). Annals of the Entomological Society of America 76: 455–462. 92. Paskewitz, S. M., and J. E. McPherson. 1983. Life history and laboratory rearing of Arhyssus lateralis (Hemiptera: Rhopalidae) with descriptions of immature stages. Annals of the Entomological Society of America 76: 477–482. 93. McPherson, J. E., B. C. Weber, and T. J. Henry. 1983. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 7. Miridae. The Great Lakes Entomologist 16: 35–42. 94. McPherson, J. E., T. E. Vogt, and S. M. Paskewitz. 1983. Effects of various split developmental photophases and constant light during each 24-hour period on adult morphology in Thyanta calceata (Hemiptera: Pentatomidae). The Great Lakes Entomologist 16: 43–46. 2012 THE GREAT LAKES ENTOMOLOGIST 119

95. Weber, B. C., and J. E. McPherson. 1983. World list of host plants of Xylosandrus germanus (Blandford) (Coleoptera: Scolytidae). Coleopterists Bulletin 37: 114–134. 96. Weber, B. C., and J. E. McPherson. 1984. The ambrosia fungus of Xylosandrus germanus (Coleoptera: Scolytidae). The Canadian Entomologist 116: 281–283. 97. McPherson, J. E., and S. M. Paskewitz. 1984. Life history and laboratory rearing of Euschistus ictericus (Hemiptera: Pentatomidae), with descriptions of immature stages. Journal of the New York Entomological Society 92: 53–60. 98. McPherson, J. E., and S. M. Paskewitz. 1984. Laboratory rearing of Amaurochrous cinctipes (Hemiptera:Pentatomidae: Podopinae) with descriptions of immature stages. Journal of the New York Entomological Society 92: 61–68. 99. Weber, B. C., and J. E. McPherson. 1984. Attack on black walnut trees by the ambrosia beetle Xylosandrus germanus (Coleoptera: Scolytidae). Forest Science 30: 846–870. 100. Vogt, T. E., and J. E. McPherson. 1985. State records and confirmations of Odonata from Illinois and Missouri. The Great Lakes Entomologist 18: 7–13. 101. Weber, B. C., and J. E. McPherson. 1985. Relation between attack by Xylosandrus germanus (Coleoptera: Scolytidae) and disease symptoms in black walnut. The Canadian Entomologist 117: 1275–1277. 102. Vogt, T. E., and J. E. McPherson. 1986. A list of the Odonata of the LaRue-Pine Hills Ecological Area. The Great Lakes Entomologist 19: 1–5. 103. McPherson, J. E., and R. J. Packauskas. 1986. Life history and laboratory rearing of Belostoma lutarium (Heteroptera: Belostomatidae) with descriptions of immature stages. Journal of the New York Entomological Society 94: 154–162. 104. Packauskas, R. J., and J. E. McPherson. 1986. Life history and laboratory rearing of Ranatra fusca (Hemiptera: Nepidae) with descriptions of immature stages. Annals of the Entomological Society of America 79: 566–571. 105. McPherson, J. E. 1986. Life history of Neoplea striola (Hemiptera: Pleidae). The Great Lakes Entomologist 19: 217–220. 106. Vogt, T. E., and J. E. McPherson. 1986. Life history and laboratory rearing of Corythucha juglandis (Hemiptera: Tingidae) with descriptions of immature stages. The Great Lakes Entomologist 19: 221–233. 107. Vogt, T. E., J. E. McPherson, and W. T. McDowell. 1986. Odonata at a small woodland pond in southern Illinois. The Great Lakes Entomologist 19: 235–237. 108. McPherson, J. E., R. J. Packauskas, and P. P. Korch, III. 1987. Life history and laboratory rearing of Pelocoris femoratus (Hemiptera: Naucoridae), with descriptions of immature stages. Proceedings of the Entomological Society of Washington 89: 288–295. 109. McPherson, J. E., and R. J. Packauskas. 1987. Life history and laboratory rearing of Nepa apiculata (Hemiptera: Nepidae) with descriptions of immature stages. An- nals of the Entomological Society of America 80: 680–685. 110. Korch, P. P., III, and J. E. McPherson. 1987. Life history and laboratory rearing of Gerris argenticollis (Hemiptera: Gerridae) with descriptions of immature stages. The Great Lakes Entomologist 20: 193–204. 111. McPherson, J. E. 1988. Life history of Mesovelia mulsanti (Hemiptera: Mesoveliidae) in southern Illinois. The Great Lakes Entomologist 21: 19–23. 112. McPherson, J. E. 1988. The need for updating life history studies. Heteropterists Newsletter 5: 11–13. 113. Keffer, S. L., and J. E. McPherson. 1988. Descriptions of nymphal instars of Abedus breviceps (Hemiptera: Belostomatidae). The Great Lakes Entomologist 21: 169–174. 114. Taylor, S. J., and J. E. McPherson. 1989. State records and confirmations of Ar- kansas flat bugs (Heteroptera: Aradidae). The Great Lakes Entomologist 22: 19–23. 120 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

115. Taylor, S. J., and J. E. McPherson. 1989. Distributional records of some Florida Aradidae (Heteroptera). Florida Entomologist 72: 265–267. 116. McPherson, J. E., and R. W. Sites. 1989. Annotated records of species of Pentato- moidea (Hemiptera) collected at lights. The Great Lakes Entomologist 22: 95–98. 117. Keffer, S. L., J. T. Polhemus, and J. E. McPherson. 1989. Notes on critical character states of Telmatotrephes (Heteroptera: Nepidae). Florida Entomologist 72: 626–629. 118. McPherson, J. E. 1989. An overview of the Hemiptera of Illinois. The Great Lakes Entomologist 22: 177–198. 119. Ahmad, I., and J. E. McPherson. 1990. Male genitalia of the type species of Corime- laena White, Galgupha Amyot and Serville, and Cydnoides Malloch (Hemiptera: Cydnidae: Thyreocorinae) and their bearing on classification. Annals of the Ento- mological Society of America 83: 162–170. 120. Keffer, S. L., J. T. Polhemus, and J. E. McPherson. 1990. What is Nepa hoffmanni (Heteroptera: Nepidae)? Male genitalia hold the answer, and delimit species groups. Journal of the New York Entomological Society 98: 154–162. 121. McPherson, J. E., R. J. Packauskas, S. J. Taylor, and M. F. O’Brien. 1990. Eastern range extension of Leptoglossus occidentalis with a key to Leptoglossus species of America north of Mexico (Heteroptera: Coreidae). The Great Lakes Entomologist 23: 99–104. 122. McPherson, J. E., and B. C. Weber. 1990. Seasonal flight patterns of Hemiptera (excluding Miridae) in a southern Illinois black walnut plantation. The Great Lakes Entomologist 23: 105–120. 123. Weber, B. C., and J. E. McPherson. 1991. Seasonal flight patterns of Scolytidae in a North Carolina and an Illinois black walnut plantation. Coleopterists Bulletin 45: 45–56. 124. McPherson, J. E. 1991. Noteworthy range extensions of three emesine species (Heteroptera:Reduviidae). The Great Lakes Entomologist 24: 99–101. 125. McPherson, J. E., S. L. Keffer, and S. J. Taylor. 1991. Taxonomic status of Melan- olestes picipes and M. abdominalis (Heteroptera: Reduviidae). Florida Entomologist 74: 396–403. 126. McPherson, J. E. 1991. Range extensions of three emesine species in North America (Heteroptera: Reduviidae). The Great Lakes Entomologist 24: 263–264. 127. McPherson, J. E. 1992. The assassin bugs of Michigan (Heteroptera: Reduviidae). Great Lakes Entomologist 25: 25–31. 128. McPherson, J. E., S. J. Taylor, and S. L. Keffer. 1992. Evaluation of characters to distinguish Fitchia aptera and F. spinosula (Heteroptera: Reduviidae). Florida Entomologist 75: 222–230. 129. Davidova-Vilimova, J., and J. E. McPherson. 1992. Pygophores of selected species of Pentatomoidea (Heteroptera) from Illinois. Acta Universitatis Carolinae Biologica 35: 143–183. 130. McPherson, J. E. 1992. [Review of] Dolling, W. R. The Hemiptera. American Ento- mologist 38: 183, 189. 131. Drickamer, L. C., and J. E. McPherson. 1993. Comparative aspects of mating behavior patterns in six species of stink bugs (Heteroptera: Pentatomidae). The Great Lakes Entomologist 25[1992]: 287–295. 132. Keffer, S. L,. and J. E. McPherson. 1993. Curictan copulation and water scorpion higher classification (Heteroptera: Nepidae). Proceedings of the Entomological Society of Washington 95: 74–78. 133. McPherson, J. E., B. C. Weber, and T. J. Henry. 1993. Seasonal flight patterns of Miridae (Hemiptera) in a southern Illinois black walnut plantation. The Great Lakes Entomologist 26: 97–106. 2012 THE GREAT LAKES ENTOMOLOGIST 121

134. McPherson, J. E. 1993. Editor’s notes. American Entomologist 39: 198–199. 135. McPherson, J. E., and S. J. Taylor. 1993. Distinguishing nymphal instars of Mesovelia mulsanti (Heteroptera: Mesoveliidae). The Great Lakes Entomologist 26: 233–236. 136. Keffer, S. L., S. J. Taylor, and J. E. McPherson. 1994. Laboratory rearing and descriptions of immature stages of Curicta scorpio (Heteroptera: Nepidae). Annals of the Entomological Society of America 87: 17–26. 137. McPherson, J. E. [Review of] Akre, R. D., G. S. Paulson, and E. P. Catts. 1994. Insects did it first. Ye Galleon Press, Fairfield, Washington. 160 pp.” American Entomologist 40: 58. 138. McPherson, J. E. [Review of] McGavin, G. C. 1994. Bugs of the world. Facts on File, Inc., New York. 192 pp. American Entomologist 40: 58–59. 139. Munyaneza, J., and J. E. McPherson. 1994. Comparative study of life histories, laboratory rearing, and immature stages of Euschistus servus and Euschistus variolarius (Hemiptera: Pentatomidae). The Great Lakes Entomologist 26: 263–274. 140. Brown, L. N., and J. E. McPherson. 1994. Life history and laboratory rearing of Gelastocoris oculatus oculatus (Fabricius) (Hemiptera: Gelastocoridae) with descriptions of immature stages. Proceedings of the Entomological Society of Washington 96: 516–526. 141. Davidova-Vilimova, J., and J. E. McPherson. 1995. History of the higher classification of the subfamily Podopinae (Heteroptera: Pentatomidae) - an historical review. Acta Universitatis Carolinae Biologica 38[1994]: 99–124. 142. Wood, D. L., and J. E. McPherson. 1995. Life history and laboratory rearing of Hydrometra hungerfordi Torre-Bueno (Heteroptera: Hydrometridae) with descriptions of immature stages. Proceedings of the Entomological Society of Washington 97: 717–728. 143. McPherson, J. E., R. C. Froeschner, and R. W. Sites. 1995. Range extension of Diaditus tejanus (Heteroptera: Reduviidae). Florida Entomologist 78: 611–612. 144. Davidova-Vilimova, S. J. Taylor, and J. E. McPherson. 1996. A new species of Mezira (Heteroptera: Aradidae) from Florida, with a key to the small Mezira of America north of Mexico. Proceedings of the Entomological Society of Washington 98: 630–639. 145. Bundy, C. S., and J. E. McPherson. 1997. Life history and laboratory rearing of Corimelaena obscura (Heteroptera: Thyreocoridae) with descriptions of immature stages. Annals of the Entomological Society of America 90: 20–27. 146. Loya, L. J., and J. E. McPherson. 1997. Life history and laboratory rearing of Oedancala dorsalis (Say) (Heteroptera: Lygaeidae) with descriptions of immature stages. Proceedings of the Entomological Society of Washington 99: 89–100. 147. McPherson, J. E., and D. L. Tecic. 1997. Notes on the life histories of Acrosternum hilare and Cosmopepla bimaculata (Heteroptera: Pentatomidae) in southern Illinois. The Great Lakes Entomologist 30: 79–84. 148. Taylor, S. J., and J. E. McPherson. 1998. Laboratory rearing of Mesovelia cryptophila (Heteroptera: Mesoveliidae). Entomological News 109: 95–98. 149. Taylor, S. J., and J. E. McPherson. 1998. Voltinism in Neogerris hesione (Heterop- tera: Gerridae) in southern Illinois. Entomological News 109: 233–239. 150. Ahmad, I., and J. E. McPherson. 1998. Additional information on male and female genitalia of Parabrochymena Larivière and Brochymena Amyot & Serville (Hemip- tera: Pentatomidae). Annals of the Entomological Society of America 91: 800–807. 151. Taylor, S. J., and J. E. McPherson. 1998. Voltinism in Merragata brunnea (Heterop- tera: Gerromorpha: Hebridae) in southern Illinois. Florida Entomologist 81: 509–515. 152. Taylor, S. J., and J. E. McPherson. 1999. Morphological variation and polyvoltinism of Microvelia pulchella (Heteroptera: Veliidae) in southern Illinois, USA. Acta Soci- etatis. Zoologicae Bohemicae 63: 237–249. 122 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

153. Hagerty, A. M., and J. E. McPherson. 2000. Survey of the Reduviidae (Heteroptera) of southern Illinois, excluding the Phymatinae, with notes on biology. The Great Lakes Entomologist 32[1999]: 133–160. 154. Hagerty, A. M., and J. E. McPherson. 2000. Life history and laboratory rearing of Arilus cristatus (Heteroptera: Reduviidae) in southern Illinois. Florida Entomolo- gist 83: 58–63. 155. Panizzi, A. R., J. E. McPherson, D. G. James, M. Javahery, and R. M. McPherson. 2000. Stink bugs (Pentatomidae), pp. 421–474. In C. W. Schaefer and A. R. Panizzi (eds.). Heteroptera of economic importance. CRC Press, Boca Raton, FL. 828 pp. 156. Taylor, S. J., and J. E. McPherson. 2000. Comparison of two population sampling methods used in field life history studies of Mesovelia mulsanti (Heteroptera: Gerromorpha: Mesoveliidae) in southern Illinois. The Great Lakes Entomologist 33[2002]: 223–230. 157. McPherson, J. E., and R. M. McPherson. 2000. Stink Bugs of Economic Importance in America North of Mexico. CRC Press, Boca Raton, FL. 253 pp. 158. Hagerty, A. M., J. E. McPherson, and J. D. Bradshaw. 2001. Life history and laboratory rearing of Emesaya b. brevipennis (Heteroptera: Reduviidae) in southern Illinois. Florida Entomologist 84: 410–414. 159. Bradshaw, J. D., and J. E. McPherson. 2001. Discovery of macroptery in Pseudo- metapterus umbrosus (Heteroptera: Reduviidae). Florida Entomologist 84: 719–720. 160. Bradshaw, J. D., and J. E. McPherson. 2002. Life history and laboratory rearing of Pseudometapterus umbrosus (Heteroptera: Reduviidae) with descriptions of immature stages. Annals of the Entomological Society of America 95: 192–200. 161. Shurtz, R. A., and J. E. McPherson. 2002. Laboratory rearing of Rocconota an- nulicornis (Heteroptera: Reduviidae) with brief notes on field life cycle. The Great Lakes Entomologist 35[2003]: 171–172. 162. Voss, S. C., and J. E. McPherson. 2003. Life history and laboratory rearing of Sinea diadema (Heteroptera: Reduviidae) with descriptions of immature stages. Annals of the Entomological Society of America 96: 776–792. 163. Taylor, S. J., and J. E. McPherson. 2003. Voltinism and laboratory rearing of Mi- crovelia hinei (Heteroptera: Gerromorpha: Veliidae). The Great Lakes Entomologist 36[2004]: 1–9. 164. Tecic, D. L., and J. E. McPherson. 2005. Resurvey of the Pentatomoidea (Heter- optera) of the La Rue-Pine Hills Research Natural Area in Union County, Illinois. The Great Lakes Entomologist 37[2004]: 30–70. 165. Shurtz, R. A., and J. E. McPherson. 2005. Life history and laboratory rearing of Sinea spinipes (Heteroptera: Reduviidae) with descriptions of immature stages. An- nals of the Entomological Society of America 98: 18–36. 166. McPherson, J. E., and I. Ahmad. 2005. Further information on male genitalia of Parabrochymena Larivière and Brochymena Amyot & Serville (Hemiptera: Pen- tatomidae). Annals of the Entomological Society of America 98: 282–290. 167. Bundy, C. S., and J. E. McPherson. 2005. Morphological examination of the egg of Mecidea major (Heteroptera: Pentatomidae). Southwestern Entomologist 30: 41–45. 168. Bundy, C. S., J. E. McPherson, and P. F. Smith. 2005. Comparative laboratory rearing of Mecidea major and M. minor (Heteroptera: Pentatomidae). Journal of Entomological Science 40: 291–294. 169. McPherson, J. E., S. J. Taylor, S. L. Keffer, and J. T. Polhemus. 2005. Life history and laboratory rearing of Macrovelia hornii (Hemiptera: Macroveliidae). Entomological News 116: 217–224. 170. McPherson, J. E., R. A. Shurtz, and S. C. Voss. 2006. Identification of nymphs of midwestern species and instars of Sinea (Hemiptera: Heteroptera: Reduviidae: Harpactorinae). 2012 THE GREAT LAKES ENTOMOLOGIST 123

Annals of the Entomological Society of America 99: 755–767. 171. McPherson, J. E., and S. J. Taylor. 2006. Notes of the field life history ofSteinovelia stagnalis (Hemiptera: Veliidae) in southern Illinois, U.S.A., with a survey of the biological literature. Entomological News 117: 399–405. 172. Taylor, S. J., and J. E. McPherson. 2006. Gerromorpha (Hemiptera: Heteroptera) in southern Illinois: species assemblages and habitats. The Great Lakes Entomolo- gist 39: 1–26. (2007). 173. McPherson, J. E., and I. Ahmad. 2007. Redescriptions of Brochymena and Parabro- chymena (Hemiptera: Heteroptera: Pentatomidae), based primarily on male genitalia, with reclassification of three species and description of New World tribe (Halyini). Annals of the Entomological Society of America 100: 673–682. 174. McPherson, J. E., and I. Ahmad. 2008. Comparison of male genitalia of Murgantia histrionica, M. varicolor, and M. violacens (Hemiptera: Heteroptera: Pentatomidae). Proceedings of the Entomological Society of Washington 110: 1028–1033. 175. McPherson, J. E., I. Ahmad, and C. S. Bundy. 2009. Comparison of the male and female genitalia of Mecidea major and M. minor (Hemiptera: Pentatomidae: Pen- tatominae: Mecideini). Journal of Entomological Science 44: 164–169. 176. Bundy, C. S., and J. E. McPherson. 2009. Life history and laboratory rearing of Corimelaena incognita (Hemiptera: Heteroptera: Thyreocoridae), with descriptions of immature stages. Annals of the Entomological Society of America 102: 1068–1076. 177. McPherson, J. E., and I. Ahmad. 2011. Murgantia thomasi, a new species of stink bug from the Dominican Republic (Hemiptera: Heteroptera: Pentatomidae). Annals of the Entomological Society of America 104: 149–153. 178. Bundy, C. S., and J. E. McPherson. 2011. Life history and laboratory rearing of Me- cidea minor (Hemiptera: Heteroptera: Pentatomidae), with descriptions of immature stages. Annals of the Entomological Society of America 104: 605–612. 179. McPherson, J. E., and I. Ahmad. 2011. Parasinea, a new genus of assassin bug with description of a new species from Colombia (Hemiptera: Heteroptera: Reduviidae). Annals of the Entomological Society of America 104: 1285–1291. 180. McPherson, J. E., R. J. Packauskas, R. W. Sites, S. J. Taylor, C. S. Bundy, J. D. Brad- shaw, and P. L. Mitchell. 2011. Review of Acanthocephala (Hemiptera: Heteroptera: Coreidae) of America north of Mexico with a key to species. Zootaxa 2835: 30–40. 181. McPherson, J. E., C. S. Bundy, and M. H. Sweet, II. 2012. Value of nymphal morphology in determining identity of pregenital and genital segments in adult Corimelaena incognita (Hemiptera: Heteroptera: Thyreocoridae). Annals of the Entomological Society of America 105: 111–117. 182. McPherson, J. E., and I. Ahmad. 2012. Comparison of male genitalia of Euschistus spp. in the midwestern United States (Hemiptera: Heteroptera: Pentatomidae). Annals of the Entomological Society of America 105: 395–402. 183. McPherson, J. E. 2012. Redescription of holotype of Sinea sericea, supplemented with description of additional males and first description of females. Annals of the Entomological Society of America 105: 613–618.

Acknowledgments I thank R. W. Sites (University of Missouri, Columbia) for his invitation to present this biographical sketch for The Great Lakes Entomologist volume honoring Dr. McPherson. I am grateful to C. S. Bundy (New Mexico State Uni- versity, Las Cruces), and R. W. Sites for lending photographs of Dr. McPherson. I also thank C. S. Bundy, K. Henderson (Silver Spring, MD), R. W. Sites, and A. G. Wheeler, Jr. (Clemson University, Clemson, SC) for reading the manuscript and offering suggestions for its improvement. 124 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 J.E. McPherson – A Career of Exemplary Service and Contributions to the Entomological Society of America George G. Kennedy Scientific societies are a vital component of the infrastructure that fosters progress in science. They play a vital role in the exchange of information through the publication of scientific journals and by organizing scientific meetings and symposia at which results and ideas are shared and discussed, professional contacts are made, and networks established and maintained. Scientific societies provide a plethora of professional development opportunities for both young and established scientists. They are volunteer organizations comprised of and led by practicing scientists, teachers, and practitioners who provide the vision, leadership, governance, scientific and ethical standards, scientific programming, and educational training opportunities that lie at the core of their mission. Exceptional service to scientific societies is often a hallmark of outstanding scientists and educators. It is therefore no surprise that outstanding service to the Entomological Society of America (ESA) spanning over 30 years is a hallmark of J. E. (Jay) McPherson’s career (Fig. 1). Recognizing the particularly important role played by the ESA Branches in the professional development of students and young scientists, Jay was active in his service to the North Central Branch, having served as vice chair of the Systematics, Morphology and Evolution Section at the Annual Meetings in 1983; as chair in 1974, 1984, and 1991; and as co-moderator for the Systemat- ics, Morphology and Evolution Section in 1994. In addition, Jay served on the branch’s Membership Committee (1980-83), Resolution Committee (1996-97) and C.V. Riley Award Selection Committee (1997-2000). Jay’s service at the ESA national level has included multiple years as a judge in the student competition for the President’s Prize, and as a member of numerous committees including the Committee on Common Names of Insects, Finance Committee, Fellow Review and Recommendation Committee, and a presidential special committee to revise qualifications for selection of ESA Fel- lows. He also served as a judge for the Linnaean Games at both the branch and national meetings over many years. However, Jay’s most significant contributions were made as a member of the editorial board and editor of American Entomologist, ESA Governing Board representative, and as ESA President during 2002. One of Jay’s most lasting contributions to the ESA was his service on the editorial board of the Bulletin of the Entomological Society of America during its transition to American Entomologist. The Bulletin, which had been published since 1955, was very narrow in scope, focusing on ESA business and affairs, with only limited scientific and general interest content. In the late 1980s the ESA Governing Board authorized the creation of a new publication with an entirely new focus and format, emphasizing high quality entomological articles of broad interest. The goal was to showcase entomology and the role of insects in shaping the human condition in the context of science, society, culture, and the environment in a way that would attract both readers and authors. This new publication, American Entomologist, first published in 1990, was intended to replace the Bulletin as a member benefit and become the signature publication of the ESA. As a member (1986-1993) and chair (1986 and 1991) of the editorial board, Jay played an important role in developing, designing, and implementing American Entomologist, while working closely

1Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613 USA 2012 THE GREAT LAKES ENTOMOLOGIST 125

Figure 1. J. E. McPherson at the Entomological Society of America Annual Meeting, Reno, NV, November 2011. Past President, Honorary Member, Fellow, Linnaean Games Head Judge, Entomological Foundation Donor, Trouble Maker. 126 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 with Paul Opler--American Entomologist’s first editor. In 1991 Jay received the ESA National Service Award in recognition of his efforts in making American Entomologist a successful publication. Jay continued to serve on the American Entomologist editorial board until 1993 when he stepped up to become its editor; a position he held until 2001 when he took office as ESA President-Elect. As editor, he expanded the scope, quality, and diversity of articles, features, and columns appearing in American Entomologist, boosting its reputation for interesting and high-quality articles. As a member of the ESA Governing Board representing the former Section A (Systematics, Morphology, and Evolution) from 1994-1996 and as President- Elect (2001), President (2002) and Past-President (2003), Jay helped the So- ciety navigate a period of great change in the processes and business models for scientific publication, one of the core functions of the ESA, and a period of financial difficulty that threatened the survival of ESA as an organization. Jay assumed office as President-Elect while the Society’s financial crisis was unfold- ing. He worked closely with Past-President Sharron Quisenberry, President Larry Larson, and the entire Governing Board to devise strategies to address the crisis by restructuring the Society’s business operations. As President, Jay implemented many of these strategies, including arranging for sale of the ESA-owned national headquarters building, dramatically reducing the number of paid staff, and outsourcing many of the business operations of the Society. These changes and a restructuring of the ESA publications enterprise were central to restoring the ESA to financial solvency and re-establishing a solid, financial foundation. Jay’s record of exemplary service to the ESA was recognized in 2004 when he was elected an Honorary Member by the Society’s membership. This recognition is reserved for “those who have served the ESA for at least 20 years through significant involvement in the affairs of the Society that has reached an extraordinary level.” In 1993, Jay was awarded the Distinguished Achieve- ment Award in Teaching by the ESA North Central Branch, and in 1996-97 he was named Outstanding Teacher in the College of Science by Southern Illinois University. In 2007 Jay was named a Fellow of the Entomological Society of America in recognition of his outstanding contributions to entomology research and teaching. Throughout his career, excellence in research and teaching, and exemplary service have been the hallmark of Jay McPherson. 2012 THE GREAT LAKES ENTOMOLOGIST 127 Mcphersonarcys, a new genus for Pentatoma aequalis Say (Heteroptera: Pentatomidae) Donald B. Thomas1

Abstract A new monotypic genus, Mcphersonarcys is erected to hold Pentatoma aequalis Say, a species formerly placed in the genus Hymenarcys. Based on the distribution of 19 character-states Hymenarcys forms a clade with its sister genus Coenus. Both genera are related to the large genus Euschistus. Mcpher- sonarcys is basal to Euschistus, whereas the clade with Hymenarcys and Coenus is derived. Removal of P. aequalis to a new genus resolves the paraphyly of Hymenarcys sensu lato.

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The genus Hymenarcys Amyot and Serville, 1843, containing four species, was revised by Rolston (1973). Regarding relationships among these Rolston stated, “The evidence seems overwhelming that H. nervosa, H. reticulata, and H. crassa are closely related species and that H. aequalis is phylogenetically distant. Erection of a monotypic genus or subgenus for H. aequalis is justifiable, but ... no practical benefit would result from a division of the genus.” It is a simple truth that in any genus containing three or more species some will be relatively close to one another and others relatively distant, thus forming objectively recognizable clades. The formal application of names to each and every such clade would tend to diminish the value of higher classifications. In this case, while I agree completely with Rolston’s analysis of relationships within Hymenarcys, his understandable restraint overlooked a deeper problem. The problem is that the suite of character states encompassed within Hymenarcys as then constituted, are inextricably entangled with those of its sister genus Coenus Dallas. Coenus was revised by Rider (1995) and contains three closely related species. Based on the characteristics that define and separate the two genera, the species described as Pentatoma aequalis Say, is so distant from its congeners that it could as well be placed in Coenus, or for that matter, Euschistus Dallas, 1851, as in Hymenarcys. Thus, one could erect a new genus, or sink all of the species into a single genus. But the present arrangement is untenable. Hymenarcys vs. Coenus. The two genera have always been difficult to separate, primarily because of H. aequalis. In Torre-Bueno’s (1939) key to the North American Pentatomini, Coenus and Hymenarcys were separated in the very last couplet. The couplet reads: “Apical part of the scutellum narrower than hemelytra ...... Hymenarcys Apical part of scutellum broader than hemelytra ...... Coenus. ” However, the widest part of the hemelytral corium is wider than the post-frenal scutellum in all species of both genera. Torre-Bueno’s error is not mitigated because he considered Coenus to be monotypic having overlooked Coe- nus inermis Harris and Johnston, a rare species described in an obscure journal

1USDA- Agricultural Research Service, Cattle Fever Tick Research Lab, 22675 N. Moorefield Road, Edinburg, TX 78541. 128 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

(Harris and Johnston 1936); or by the subsequent discovery and description of Coenus explanatus Rider. McPherson’s (1982) key to the genera of northeastern North America has Coenus and Hymenarcys separating at couplet 9 which reads, “Hind tibiae distinctly sulcate dorsally throughout their lengths..... Coenus Hind tibiae not distinctly sulcate throughout their lengths....Hymenarcys. ” But according to Rolston and McDonald (1984) the hind tibiae are sulcate in both genera which belies the fact that the species of Hymenarcys vary in the degree of sulcation with H. crassa Uhler being mostly asulcate. In their key to the genera of the western Hemisphere Pentatomini Part III, Hymenarcys keys out in two places, with H. aequalis falling out separately, scored as having the bucculae evanescent (vs. lobed) posteriorly. But the bucculae, while not clearly lobate, are strongly truncate in H. aequalis and certainly not evanescent or arcuate. Hymenarcys can be separated from Coenus because in the former the scutellum is distinctly shorter than the hemelytral corium. Thus, while Rolston and McDonald retained the two traditional taxa, their dichotomous arrangement based on two characters, separated the included species into three natural groups. Consideration of a larger suite of characters gives a similar result. Character States. Males and females of all seven included species were examined. Rider (1995) states that Coenus belongs to a group of genera which includes Euschistus and that it is most closely related to Hymenarcys. Rolston (1974) provided a key including Hymenarcys for those genera related to Euschistus Dallas. Therefore the nominate subgenus of Euschistus, type- species Euschistus tristigmus (Say), is used as the out-group for the purpose of establishing the polarity of character states. A total of 19 characters were found to have variable states among these species. Male Genitalic characters. 1. Dorsal Thecal Appendages. All species of Hymenarcys, all species of Coenus and all species of Euschistus have these appendages. According to Rider (1995) this character is unique to the genera related to Euschistus and is thus an important synapomorphy indicating a common ancestry for the taxa under consideration. 2. Thecal Lobes. In all species of Coenus, Euschistus tristigmus, most but not all species of the subgenus Euschistus, and all species of Hymenarcys except H. aequalis, the theca has lateral lobes. All species of Coenus and all species of Hymenarcys except H. aequalis have a well developed dorsal lobe. The dorsal lobe is absent in E. tristigmus and most but not all species of the subgenus Euschis- tus. In H. aequalis the sides and dorsum of the theca is simple, devoid of lobes. 3. Distiphallus. In all species of Coenus, all species in the nominate subgenus Euschistus (except E. schaffneri Rolston and E. zafadus Rolston) and all species of Hymenarcys except H. aequalis, the distiphallus (penisfilum) is long and coiled. In H. aequalis the distiphallus (illustrated by Rolston 1973) is about as long as the theca but not long enough to coil. 4. Inferior Ridge of Pygophore. All species of Euschistus and all species of Coenus have a ridge just ental to the posterior margin of the pygophore. None of the species of Hymenarcys have an inferior ridge except H. aequalis (Fig. 1). 5. Proctigeral tubercles. In all species of the nominate subgenus Euschis- tus, all species of Coenus and all species of Hymenarcys, except H aequalis, the tubercles on the proctiger are located subapically. In H. aequalis the tubercles are submedial (Fig. 1). Rolston (1974) accorded this character some importance in defining subgenera ofEuschistus . 6. Pygophoral Tooth. Coenus delius (Say) is unique among all of the included species in having an erect tooth on the posterior margin of the pygophore. 2012 THE GREAT LAKES ENTOMOLOGIST 129

Figure 1. Male pygophore, dorsal view. Proc. T = proctigeral tubercle. Inf. R. = inferior ridge.

Figure 2. Female terminalia, postero-ventral view. 130 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Female genitalic characters. 7. Spiracles. The eighth paratergite (Fig. 2) of all species of Euschistus (except E. zafadus), Coenus and Hymenarcys lack a functional spiracle. The spiracle is present in most Pentatomine genera including some of those considered to be related to Euschistus. It is unlikely in the extreme that the spiracle would re-evolve after being lost and thus the absence of the spiracle is the apomorphic condition. 8. Spermathecal Duct. The spermathecal duct is relatively long and convo- luted in all species of Coenus and all species of Hymenarcys except H. aequalis. In H. aequalis and in Euschistus the spermathecal duct is short and simple. Somatic characters. 9. Scutellar fovea. In all species of Euschistus, all species of Coenus, and all species of Hymenarcys except H. nervosa (Say), the fovea are lacking. Hymenar- cys nervosa is unique in having a foveum at each basal angle of the scutellum. 10. Paraclypeal length. Hymenarcys crassa is unique in having the paraclypei much longer than the clypeus. In all other species of Hymenarcys, all species of Coenus and Euschistus tristigmus, and most species of Euschistus (including all of those in the nominate subgenus), the clypeus and paraclypei are equal or subequal in length. 11. Buccular terminus. In all species of Euschistus, including E. tristigmus, the bucculae are evanescent posteriorly. In H. aequalis the terminus of the bucculae is truncate. In all species of Coenus and all other species of Hymenarcys, the terminus of the bucculae is lobate. Rolston and McDonald (1984) mistakenly included H. aequalis in a branch of their key with those pentatomine genera that have the buccular terminus evanescent. 12. Rostral Length. In all species of Hymenarcys except H. aequalis, the apex of the rostrum does not attain the metacoxae. In all species of Coenus and Euschistus, and H. aequalis the apex of the rostrum attains the posterior margin of the metacoxae. 13. Clypeal punctation. In all species of Coenus the clypeus is contrastingly devoid of punctures. In all species of Hymenarcys and in most Euschistus including E. tristigmus, the clypeus has many punctures, typically as dense as on the paraclypei. 14. Hemelytral membrane. In all species of Coenus and all species of Hymenarcys except H. aequalis, the veins of the membrane are reticulate. In H. aequalis and in all species of the nominate subgenus of Euschistus (except E. schaffneri) the venation is simple. 15. Anterolateral pronotal margin. Species of Euschistus characteristically have the margin dentate. In Hymenarcys and Coenus the margin is edentate. 16. Pronotal shape. Hymenarcys nervosa is unique in having the pronotal margin convex in dorsal view. All other species of Hymenarcys, all species of Coenus, and in all species in the nominate subgenus of Euschistus (except E. schaffneri), the margin is subrectilinear or even somewhat concave (e.g., Coenus explanatus). 17. Punctation of pronotal margin. In Hymenarcys crassa and in the species of Coenus, the inframargin of the anterolateral pronotal margin is contrastingly devoid of punctures. In the other species of Hymenarcys and in Euschistus tristigmus, the punctations of the disc extend to the edge of the pronotum. 18. Hemelytral corium. In all species of Hymenarcys and Euschistus the hemelytral corium extends distinctly past the apex of the scutellum. In all species of Coenus the hemelytral corium does not distinctly surpass the apex of the corium. 2012 THE GREAT LAKES ENTOMOLOGIST 131 19. Metasternum. In all species of Hymenarcys except H. aequalis the metasternum is sulcate. In all species of Coenus, Euschistus and H. aequalis the metasternum is flat. Character Analysis. Of the 19 included characters two had character states common to all species of Euschistus, Coenus, and Hymenarcys sensu Rolston: the spiracles vestigial on the female 8th paratergite and the presence of thecal appendages in the males. This synapomorphy is significant as evidence of common ancestry of the included species. Six of the characters had states unique to single species: the long rostrum in H. aequalis, the truncate buccular terminus in H. aequalis, the scutellar fovea of H. nervosa, the convex anterolateral pronotal margin of H. nervosa, the medial pygophoral tooth of Coenus delius and the elongated paraclypei in H. crassa. Because these states were unique at the species level they provide no information on relationships. It is the pattern of shared characteristics that provide evidence of the latter. All species of Euschistus have the anterolateral pronotal margins dentate and the terminus of the bucculae evanescent. None of the species of Coenus or Hymenarcys sensu Rolston have either of these two character states. Thus, Euschistus forms a clade separable from the other included taxa. In all species of Coenus the clypeus is impunctate and the corium fails to surpass the apex of the scutellum whereas in all species of Euschistus and Hymenarcys the corium is distinctly longer than the scutellum and the clypeus is densely punctate. Thus Coenus as constituted in the revision by Rider forms a clade separable from Euschistus and Hymenarcys sensu Rolston. However, there are no synapomorphic character states that unite the species of Hymenarcys unless one eliminates H. aequalis and thus Hymenar- cys sensu Rolston is paraphyletic. But when H. aequalis is removed the genus Hymenarcys sensu novo can be defined by having a sulcate metasternum (in H. aequalis, Coenus and Euschistus the metasternum is flat) and by having a pygophore without an inferior ridge (all species of Euschistus, Coenus and H. aequalis have an inferior ridge). Thus Hymenarcys sensu novo is a definable clade. Furthermore, the genera Coenus and Hymenarcys sensu novo form a clade separable from Euschistus and H. aequalis because they share the character states of having the spermathecal duct long and convolute (vs. short and simple in the latter) and the terminus of the bucculae lobate (vs. evanescent in Euschistus and truncate in H. aequalis). Further support for this clade is found in the hemelytral membrane which is reticulate in all Coenus and Hymenarcys sensu novo but simple in H. aequalis and in the subgenus Euschistus (except E. schaffneri). Lastly, Coenus, Hymenarcys sensu novo and and almost all species of Eus- chistus form a clade separable from H. aequalis because they have a long, coiled distiphallus (short and simple in H. aequalis) and the theca of the aedeagus lobate (entire in H. aequalis). Thus, H. aequalis lacking the synapomorphies found even in most Euschistus, is not just an aberrant species, but clearly not even related to either Hymenarcys or to Coenus. In a genus as large as Euschistus some secondarily evolved plesiomorphies can be expected. But the aforementioned aberrant species E. zafadus and E. schaffneri have combinations of plesiomorphic characters that lead this author to believe that they are wrongly placed in the subgenus Euschistus, yet, they lack the synapomorphies which define the other subgenera. Their reassignment must be held in abeyance until further study. In any case a new genus is required to hold H. aequalis and I here propose the name Mcphersonarcys. Mcphersonarcys Thomas, new genus (Fig. 3). Type-species: Pentatoma aequalis Say, 1832, by monotypy and herein designated. 132 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Figure 3. Mcphersonarcys aequalis (Say), dorsal habitus. 2012 THE GREAT LAKES ENTOMOLOGIST 133 Description: Head length subequal to pronotal length at midline; eyes subpedunculate. Anterolateral pronotal margin carinate, simple, not dentate, not reflexed; humeral angles not produced; anterolateral angles with a tooth. Width of scutellum at end of frenum slightly greater than half basal width. Hemelytral corium longer than scutellum, terminating at fourth visible connexival segment; hemelytral membrane with simple venation. Abdominal venter without basal tubercle or spine. Metasternum flat. Metathoracic scent gland orifice with short auriculate peritreme. Femora unarmed; tibiae shallowly sulcate. Clypeus and paraclypei subequal in length, apically obtuse; basal antennal segment not quite reaching apex of head. Bucculae truncate at posterior terminus; apex of first rostral segment not surpassing posterior terminus of bucculae. Eighth pa- ratergites of female lacking spiracles; spermathecal duct short and simple. Male pygophore with a transverse ridge seated ental to the posterior margin (inferior ridge); tubercles on dorsum of proctiger submedial in position; distiphallus of aedeagus not longer than the theca; theca simple, without lobes. Etymology: Derived from a combination of the surname McPherson and the suffix of the name Hymenarcys, the genus to which the only species was formerly combined. The new genus is named to honor Prof. Jay McPherson of Southern Illinois University for his contributions to the knowledge of the family of which this genus is a member. Material Examined: Kansas: Onaga: Crevecceur. Riley: Manhattan. Brown: Hiawatha. Mississippi: Sunflower: Indianola. Missouri: Boone: Co- lumbia. Bloomsdale: Bloomsdale. Texas: Kerr: Kerrville.

Acknowledgments The habitus illustration was prepared by Robert Sites (University of Missouri). Joseph Eger (Dow AgroSciences) and David Rider (North Dakota State University) provided reviews of the manuscript. Mention of a product does not constitute endorsement by the USDA, an equal opportunity provider and employer.

Literature Cited Harris, H. M., and H. G. Johnston. 1936. A new genus and species of Podopidae and a new Coenus (Hemiptera: Scutelleroideae [sic]). Iowa State College Journal of Sci- ence 10: 377–380. McPherson, J. E. 1982. The Pentatomoidea (Hemiptera) of northeastern North America with emphasis on the fauna of Illinois. Southern Illinois University Press, Carbondale IL. 240 pp. Rider, D. A. 1995. Review of the genus Coenus Dallas, with the description of C. explanatus, new species (Heteroptera: Pentatomidae). Journal of the New York Entomological Society 103: 39–47. Rolston, L. H. 1973. A review of Hymenarcys (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 81: 111–117. Rolston, L. H. 1974. Revision of the genus Euschistus in middle America (Hemiptera, Pentatomidae, Pentatomini). Entomologica Americana 48: 1–102. Rolston, L. H., and F. J. D. McDonald. 1984. A conspectus of Pentatomini of the Western Hemisphere, Part 3. (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 92: 69–86. Torre-Bueno, J. R. de la. 1939. A synopsis of the Hemiptera-Heteroptera of America north of Mexico. Part I. Families Scutelleridae, Cydnidae, Pentatomidae, Aradidae, Dysodiidae and Termitaphididae. Entomologica Americana 19: 141–304. 134 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 The Stink Bugs (Hemiptera: Heteroptera: Pentatomidae) of Missouri Robert W. Sites1,2, Kristin B. Simpson2, and Diane L. Wood3

Abstract The stink bug (Hemiptera: Pentatomidae) fauna of Missouri was last treated more than 70 years ago. Since then, many more specimens have become available for study, substantial papers on regional faunas have been published, and many revisions and other taxonomic changes have taken place. As a consequence, 40% of the names from the previous Missouri state list have changed or the taxa have been removed. The fauna of stink bugs known from Missouri based on specimens from museums, other collections, and the literature is presented. Included for each species is a general overview of biological and ecological information, overall distribution, and local distribution among Mis- souri Natural Divisions. Also presented for Missouri specimens are biological label data and extreme dates of adult collection. All totaled, 57 species are now known from the state, eight of which represent new state records: Asopinae: Alcaeorrhynchus grandis (Dallas), Tylospilus acutissimus (Stål); Pentatominae: Banasa calva (Say), Banasa euchlora Stål, Cosmopepla intergressa (Uhler), Halyomorpha halys (Stål), Neottiglossa undata (Say); Podopinae: Amaurochrous brevitylus Barber and Sailer.

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As a family of true bugs, the stink bugs (Heteroptera: Pentatomidae) have received a great deal of notoriety by scientists and non-scientists in part because their common name engenders curiosity; they are large, apparent and sometimes brightly colored; some are crop pests and others are predators of crop pests. This is one of the largest families of Heteroptera and includes species exhibiting a wide variety of biological and ecological attributes. Although most of the literature on stink bugs focuses on questions related to controlling populations of destructive species (see McPherson and McPherson 2000), a siz- able body of literature exists on the taxonomy and systematics of stink bugs at various taxonomic levels, including superfamily, family, and genus. On a broad taxonomic and geographic scale, the Pentatomoidea of northeastern North America was presented by McPherson (1982) with keys, distributions, and exhaustive reviews of biology. The stink bug fauna of many individual states with a similar species composition to that of Missouri have been presented over the past century, including for Arkansas (Barton and Lee 1981), Illinois (Hart 1919, McPherson 1979b), Iowa (Stoner 1920), Kansas (Gates and Peters 1962), Michigan (McPherson 1970, 1979a), Nebraska (Zimmer 1911), Ohio (Furth 1974), and Virginia (Hoffman 1971). The fauna of Pentatomoidea of Missouri was presented by Froeschner (1941) with an annotated list and taxonomic key to species. However, that paper is more than 70 years old and since then, many additional specimens have become available for study in

1Corresponding author (email: [email protected]). 2Enns Entomology Museum, Division of Plant Sciences, University of Missouri, Colum- bia, Missouri 65211, U.S.A. 3Department of Biology, Southeast Missouri State University, Cape Girardeau, Mis- souri 63701, U.S.A. (email: [email protected]). 2012 THE GREAT LAKES ENTOMOLOGIST 135 museum and other collections, other papers addressing regional faunas have been published, and many revisions and other taxonomic changes have taken place. As a consequence, 40% of the names from the 1941 Missouri state list have changed and various taxa have been removed from or added to the list. Thus, presented here is an annotated list of the stink bugs currently known from Missouri based on museum and other collection records, and the literature.

Materials and Methods Data were gathered from the Enns Entomology Museum - University of Missouri, other collections, a state agency, and the literature. In the Distri- bution sections, we provide an overall distribution of each species followed by distribution in Missouri as associated with six Natural Divisions (Thom and Wilson 1980). The Natural Divisions are the result of a hybridized regionaliza- tion scheme based on an integration of physical (geologic history, soils, bedrock geology, topography), biological (known plant and animal distributions, presettle- ment vegetation), and other natural factors (Fig. 1).

Figure 1. Natural Divisions of Missouri, modified from Thom and Wilson (1980). Reproduced with permission by Richard Thom and James Wilson. 136 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

A project is in progress to digitize the Enns Entomology Museum which is expected to make available high resolution images of the specimen holdings and legible label data, including the pentatomids referenced here. Unless otherwise indicated, records presented here are from the Enns Entomology Museum (MU). Data from the Missouri Department of Agriculture are indicated as “MDA,” Southeast Missouri State University as “SEMO,” and the University of Missouri Delta Center at Portageville as “UMDC.” Because many county records referenced by Froeschner (1941) are represented by specimens in the Enns Entomology Museum, his county records are included with those of the Museum. Occurrence in St. Louis is noted separately because the city is its own geopolitical entity and has no county association. The date extremes given for each species represent the earliest and latest dates of collection of adults. Pentatomidae Known from Missouri Biology overview. In general, species of the subfamily Asopinae are predators as adults, whereas those of Pentatominae and Podopinae are phytophagous, although exceptions exist. In Missouri, species are univoltine or bivoltine (Oetting and Yonke 1971c) and adults of most species overwinter in association with some type of protection on the ground. Immature stages pass through five nymphal instars and development from egg to adult generally requires 6–8 weeks (Oetting and Yonke 1971c). Taxonomic overview. All totaled, 57 species are known from Missouri from the following three subfamilies: Asopinae (12), Pentatominae (43), and Podopinae (2). Eight of the species, representing all three subfamilies, are newly reported for the state of Missouri.

SUBFAMILY – ASOPINAE Alcaeorrhynchus grandis (Dallas) Canthecona grandis Dallas, 1851: List Hem. Brit. Mus. 1: 91. Alcaeorrhynchus grandis: Bergroth, 1891, Rev. Ent. 10: 235. Distribution. Missouri south to Florida, Texas, and Argentina. Mis- souri records are from 5 counties in Ozark and Mississippi Lowlands Natural Divisions (UMDC). Label data. Biological: alfalfa, nest of malacocana, light, light trap (UMDC). Date extremes in Missouri: 14 April–29 July (UMDC). Discussion. This species is sometimes commonly known as the giant strong-nosed stink bug (Richman and Mead 2011) and is the largest of the Asopinae in Missouri. It was reared in the laboratory by Richman and Whitcomb (1978) and has been reported as a predator of butterfly larvae (Araujo e Silva 1933). This is the first record of this species from Missouri.

Apoecilus cynicus (Say) Pentatoma cynica Say, 1831b: Descrip. Het. Hem. N. Am., p. 3. Apoecilus cynicus: Thomas, 1992, Thom. Say Found. Mon. 16: 27. Distribution. Arizona to “Dakota” east to Florida and New York (Froe- schner 1988), and Canada (Thomas 1992). Missouri records are from the City of St. Louis and 17 counties in Ozark, Glaciated Plains, and Ozark Border Natural Divisions. Label data. Biological: eggs under bark black oak (13 March), in fire- wood logs, under bark, lights. Date extremes in Missouri: 5 May–19 October, 3rd instars 11 May. 2012 THE GREAT LAKES ENTOMOLOGIST 137

Discussion. The biology of this species (as Apateticus cynicus) was reviewed by Jones and Coppel (1963) and McPherson and Mohlenbrock (1976), and biology and distribution by McPherson (1982). This large species usually is found under bark and although it was described from Missouri, it is uncommon in the state (Froeschner 1941).

Euthyrhynchus floridanus (Linnaeus) Cimex floridanus Linnaeus, 1767: Syst. Nat. Ed. 12, 1, pt. 2: 719. Euthyrhynchus floridanus: Dallas, 1851, List Hem. Brit. Mus. 1: 104. Distribution. Pennsylvania to Missouri south to Brazil (Froeschner 1988). Missouri records are from 3 counties in Ozark and Mississippi Lowlands Natural Divisions. Label data. Biological: aggregated on bee hive, honeysuckle. Date extremes in Missouri: 10 September–28 September, 3rd instars on 6 June. Discussion. This species was reported from Missouri by Oetting and Yonke (1975). It was reared in the laboratory by Richman and Whitcomb (1978). Information on biology and immature stages was provided by Oetting and Yonke (1975), and biology and distribution were reviewed by McPherson (1982).

Perillus bioculatus (Fabricius) Cimex bioculatus Fabricius, 1775: Syst. Ent., p. 715. Perillus bioculatus: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 129. Distribution. Alberta to Quebec south to Arizona, Florida, and Mexico (Froeschner 1988). Missouri records are from the City of St. Louis and 7 counties in Glaciated Plains, Ozark Border, and Big Rivers Natural Divisions. Label data. Date extremes in Missouri: 11 May–12 December (MU, Froeschner 1941). Discussion. Perillus bioculatus is known commonly as the twospotted stink bug. This predaceous species was introduced to Europe for control of the Colorado potato beetle (Thomas 1992) and was mass-reared for use against this pest (Franz 1961). A technique to rear this species on artificial diet was presented by Coudron and Kim (2004). The biology and distribution of P. bioculatus were reviewed by McPherson (1982). Adults of this species exhibit color dimorphism in which one form is red or orange and black, and the other form is white or yellow and black. The latter is the ‘clanda’ form and was described by Say (1825) from Missouri. This species was considered by Froeschner (1941) to be scarce in Missouri.

Perillus circumcinctus Stål Perillus circumcinctus, Stål 1862: Stet. Ent. Zeit. 23(1): 89. Distribution. Saskatchewan to New Hampshire south to Nebraska, Missouri, and Ohio (Froeschner 1988). Discussion. This predaceous stink bug specializes on Chrysomelidae as prey (Evans 1982, and citations therein). The biology and distribution were reviewed by McPherson (1982). Although P. circumcinctus was listed from Mis- souri by Van Duzee (1917) and Froeschner (1988), we have seen no specimens from the state and know of no specific locality data. Because this species is northern in its overall distribution and is known from only the northern half of Illinois (McPherson 1982), it should be expected in the Glaciated Plains Natural Division of Missouri. 138 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Perillus exaptus (Say) Pentatoma exapta Say, 1825: J. Acad. Nat. Sci. Phila. 4: 313. Perillus exaptus: Stål, 1870, K. Svens. Vet.-Akad. Handl. 9(1): 32. Distribution. British Columbia to California east to Nova Scotia and New Jersey (Froeschner 1988). The sole Missouri record is from St. Louis (Froeschner 1941). Label data. Biological: weedy field. Date of collection: 2 July 1935. Discussion. The biology and distribution of this species were reviewed by McPherson (1982). This species was described based on specimens from Mis- souri. We have not seen specimens of this species from Missouri, although the record of a single specimen swept from a weedy field in St. Louis was presented by Froeschner (1941) and the species was listed from Missouri by Uhler (1861), Van Duzee (1917), and Blatchley (1926).

Perillus strigipes (Herrich-Schäffer) Podisus strigipes Herrich-Schäffer, 1853: Wanz. Ins. 9: 338. Perillus strigipes: Hoffman, 1971, Va. Polyt. Inst. Res. Div. Bull. 4: 55. Distribution. Massachusetts to Florida west to Colorado and Arizona (Froeschner 1988). Missouri records are from 6 counties in Osage Plains, Glaci- ated Plains, Ozark, and Ozark Border Natural Divisions. Label data. Biological: Asclepias, Japanese beetle trap (Froeschner 1941). Date extremes in Missouri: 3 May–3 September. Discussion. This species was reported from Missouri by Froeschner (1941). The biology and distribution (as Mineus strigipes) were reviewed by McPherson (1982).

Podisus maculiventris (Say) Pentatoma maculiventris Say, 1831a: Descrip. N. Am. Ins. La., p. 11. Podisus spinosus: Stål, 1867, Öfv. K. Svens. Vet.-Akad. Förh. 24(7): 497. Podisus maculiventris: Torre-Bueno, 1903, J. N. Y. Ent. Soc. 11: 128. Distribution. British Columbia to Quebec south to California, Texas, and Florida (Froeschner 1988). Missouri records are from the City of St. Louis and 81 counties in all six Natural Divisions (MU, MDA). Label data. Biological: sucking Catocala larva, Phlox diverticula, Rhus canadensis, Salix, alfalfa, alsike, feeding on adult bibionid, corn, fescue, La- dino clover, partridge pea, pre-tassel corn, rank grass, red clover, white clover, cheat, fumigation collection soybeans, roadside sweeps, sticky board, sweeping garden, sweeping weeds, blacklight, light trap, lights, malaise, mercury vapor light (MU); sugarbeet, bollworm trap (UMDC). Date extremes in Missouri: 14 April–26 November (MU), nymphs in July–August (Froeschner 1941). Discussion. This species is known commonly as the spined soldier bug. The biology was discussed by McPherson and Mohlenbrock (1976), a list of prey species was given by McPherson (1980a), laboratory rearing data were presented by Richman and Whitcomb (1978), rearing on artificial diet by Wit- tmeyer and Coudron (2001), and biology and distribution were reviewed by McPherson (1982). In Missouri, adults have been observed preying on larvae of Lepidoptera and Lema trilineata (Olivier) (Coleoptera: Chrysomelidae) (Froeschner 1941). 2012 THE GREAT LAKES ENTOMOLOGIST 139

Podisus placidus Uhler Podisus placidus Uhler, 1870: Am. Ent. 2: 203. Distribution. British Columbia to Quebec south to Utah, Arkansas, and New Jersey (Froeschner 1988). Missouri records are from 7 counties in Ozark and Ozark Border Natural Divisions. Label data. Biological: feeding on Hyphantria cunea (Drury), hickory. Date extremes in Missouri: 20 April–12 September. Discussion. The biology and immature stages were described by Oetting and Yonke (1971a). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). Froeschner (1941) considered this species to be rare in Missouri.

Podisus serieventris Uhler Podisus serieventris Uhler, 1871: Proc. Bost. Soc. Nat. Hist. 14: 94. Distribution. British Columbia to Newfoundland south to New Mexico and North Carolina (Froeschner 1988). Missouri records are from 4 counties in Ozark, Ozark Border, and Glaciated Plains Natural Divisions. Label data. Biological: blacklight, mercury vapor light. Date extremes in Missouri: 22 July–10 October. Discussion. This species was first reported from Missouri by Froeschner (1941) based on a specimen from St. Louis County. Information on the biology of this species was given by Prebble (1933) and biology and distribution were reviewed by McPherson (1982).

Stiretrus anchorago (Fabricius) Cimex anchorago Fabricius, 1775: Syst. Ent., p. 699. Stiretrus anchorago: Dallas, 1851, List Hem. Brit. Mus. 1: 80. Distribution. Alberta to New York south to New Mexico, Florida, and Panama (Froeschner 1988). Missouri records are from the City of St. Louis and 30 counties in Ozark, Ozark Border, Glaciated Plains, Mississippi Lowlands, and Big Rivers Natural Divisions (MU, UMDC). Label data. Biological: Asclepias attacking larva, alfalfa, beans, preying on e. tent caterpillar, hibiscus, potato plant, red clover, sweeping in aban- doned field, weed, malaise trap (MU); JB [Japanese) beetle] trap (MDA). Date extremes in Missouri: 14 April–15 October (UMDC, MU); nymphs in August (Froeschner 1941). Discussion. This species is variable in its expression of color for which many forms have been named (Thomas 1992). Immature stages were described by Oetting and Yonke (1971a) and laboratory rearing data were presented by Richman and Whitcomb (1978). Various aspects of biology of this species were presented by Waddill and Shepard (1974) and McPherson and Mohlenbrock (1976), and a review of the literature by McPherson (1982). This species was considered to be relatively scarce in Missouri (Froeschner 1941).

Tylospilus acutissimus (Stål) Podisus acutissimus Stål, 1870: K. Svens. Vet.-Akad. Handl. 9(1): 53. Tylospilus acutissimus: Uhler, 1876, Bull. U. S. Geol. Geogr. Surv. Terr. 1: 283. Distribution. Colorado and Arizona south through Texas to Guatemala 140 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

(Froeschner 1988). The sole Missouri record is from Columbia, Boone County in the Ozark Border Natural Division. Label data. Date collected: 1 November 1975. Discussion. Information concerning the biology of T. acutissimus was presented by Stoner (1920). The specimen for this apparently extralimital record was determined by K. A. Phillips and confirmed by D. A. Rider. This is the first record of this species from Missouri.

SUBFAMILY – PENTATOMINAE Aelia americana Dallas Aelia americana Dallas 1851: List Hem. Brit. Mus. 1: 223. Distribution. British Columbia to Michigan south to Arizona and Alabama (Froeschner 1988). Missouri records are from 18 counties in Ozark, Ozark Border, Glaciated Plains, Mississippi Lowlands, and Big Rivers Natural Divisions. Label data. Biological: Andropogon. Date extremes in Missouri: 5 Feb- ruary–17 December. Discussion. Although this species has been considered rare (Froeschner 1941, McPherson and Mohlenbrock 1976), it has been collected commonly from apple trees in orchards and has been reported to overwinter in grass clumps and under mullein leaves (Froeschner 1941).

Banasa calva (Say) Pentatoma calva Say, 1831b: Descrip. Het. Hem. N. Am., p. 7. Banasa calva: Uhler, 1876, Bull. U. S. Geol. Geogr. Surv. Terr. 1: 291. Distribution. Maine to British Columbia south to Georgia, Mexico, and Guatemala (Froeschner 1988). Missouri records are from 4 counties in Missis- sippi Lowlands, Glaciated Plains, and Ozark Border Natural Divisions. Label data. Biological: blacklight, light trap. Date extremes in Missouri: 21 May–3 September. Discussion. Thomas and Yonke (1981) reviewed the genus. The life history of B. calva was reported by DeCoursey (1963), and the biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). This is the first record of this species from Missouri.

Banasa dimidiata (Say) Pentatoma dimiata Say, 1831b: Descrip. Het. Hem. N. Am., p. 7. Pentatoma dimidiata: LeConte, 1859, Comp. Writ. T. Say, 1: 318. Banasa dimidiata: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 43. Distribution. British Columbia to Nova Scotia south to New Mexico and Florida (Froeschner 1988). Missouri records are from 10 counties in Ozark, Ozark Border, Osage Plains, Glaciated Plains, and Mississippi Lowlands Natural Divisions (MU, UMDC). Label data. Biological:, Spirea flower, blacklight, light, mercury vapor light (MU); light trap (UMDC); Japanese beetle trap (Froeschner 1941). Date extremes in Missouri: 1 May–19 September. Discussion. Thomas and Yonke (1981) reviewed the genus and reported this species on blueberry in Missouri. Say (1831b) named this species dimiata; 2012 THE GREAT LAKES ENTOMOLOGIST 141 however, LeConte (1859) emended the specific epithet to dimidiata, which is what he thought Say had intended. Although this was an unjustified emendation, most authors have followed LeConte, and the usage of dimidiata for over 150 years, albeit inconsistent, has created some stability and justifies its continued use (see explanation in Hoffman 2005). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982).

Banasa euchlora Stål Banasa euchlora Stål, 1872: K. Svens. Vet.-Akad. Handl. 10(4): 44. Distribution. New York to Florida west to Arizona and Utah (Froeschner 1988), Canada (Paiero et al. 2003), and northern Mexico (Thomas and Yonke 1981). Missouri records are from 3 counties in Ozark, Ozark Border, and Osage Plains Natural Divisions. Label data. Biological: house basement, blacklight trap, mercury vapor light. Date extremes in Missouri: 4 May–31 December. Discussion. Thomas and Yonke (1981) reviewed the genus. This species is associated with cedar (McPherson 1982, and citations therein). The biology and distribution of B. euchlora were reviewed by McPherson (1982). This is the first record of this species from Missouri.

Banasa sordida (Uhler) Atomosira sordida Uhler, 1871: Proc. Bost. Soc. Nat. Hist. 14: 98. Banasa sordida: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 44. Distribution. Southern Canada, the contiguous U. S., and northern Mexico (Thomas and Yonke 1981). Missouri records are from 3 counties in Ozark Border, Glaciated Plains, and Mississippi Lowlands Natural Divisions (MU, UMDC). Label data. Biological: cow peas, blacklight, light trap, mercury vapor light (MU); cow peas (UMDC). Date extremes in Missouri: 6 June–18 September. Discussion. Thomas and Yonke (1981) reviewed the genus and reported this species in Missouri to use J. (Juniperus) virginiana as a host plant. The biology and distribution of B. sordida were reviewed by McPherson (1982).

Brochymena cariosa Stål Brochymena cariosa Stål, 1872: K. Svens. Vet.-Akad. Handl. 10(4): 17. Distribution. North Carolina to Florida west to Nebraska and Louisi- ana (Froeschner 1988). Missouri records are from the City of St. Louis and 19 counties in Ozark, Ozark Border, Osage Plains, Mississippi Lowlands, and Big Rivers Natural Divisions (MU, SEMO, UMDC). Label data. Date extremes in Missouri: 1 February–15 December (MDA, MU). Discussion. The biology and distribution of this species were reviewed by McPherson (1982). This species overwinters under bark and in logs and is known to congregate in large numbers (Froeschner 1941).

Brochymena carolinensis (Westwood) Cimex annulata Fabricius, 1775: Syst. Ent., p. 704. (preoccupied) Halys carolinensis Westwood, 1837: Hope. Cat. 1: 22. Brochymena carolinensis: Dallas, 1851, List Hem. Brit. Mus. 1: 189. 142 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Distribution. New York to Florida west to Alberta and Texas (Froeschner 1988). Missouri records are from 2 counties in Ozark Natural Division (MU, McPherson 1980b). Label data. Biological: at a nursery. Date extremes in Missouri: 26 August–3 September. Discussion. This species was first reported from Missouri by McPherson (1980b). Biology and distribution were reviewed by McPherson (1980b, 1982).

Brochymena punctata punctata Van Duzee Brochymena punctata Van Duzee, 1909: Can. Ent. 41: 369. Distribution. Virginia to Florida (Froeschner 1988) west to Oklahoma and Missouri (Larivière 1992). Discussion. We have not seen specimens of this species from Missouri, although Larivière (1992) indicated that a specimen with a Missouri label is in the U.S. National Museum. The species has been collected on black walnut (McPherson 1976c) and oaks (Ruckes 1946b). The biology and distribution of B. p. punctata were reviewed by McPherson (1982). Larivière (1992) considered this species to be rare.

Brochymena quadripustulata (Fabricius) Cimex 4.pustulatus Fabricius, 1775: Syst. Ent., p. 704. Brochymena serrata: Amyot and Serville, 1843, Hist. Nat. Ins. Hem., 107. Brochymena quadripustulata: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 16. Distribution. British Columbia to Quebec south to Arizona and Florida (Froeschner 1988). Missouri records are from the City of St. Louis and 49 counties in all six Natural Divisions (MU, MDA, UMDC). Label data. Biological: Cirsium sp., Fraxinus penasylvanica [sic] Marsh trunk, Helianthus, Populus deltoides Bartr. foliage, Quercus palustris trunk, elm, green ash twig, jack pine, oak & poplar, pin oak leaf, pin oak, soybeans, willow, herbage, prairie, under bark of dead oak [18 Jan.], under bark [11 Jan. and 20 Apr.], under bark of stump [10 Nov.], under stone near woods [16 Dec.] (MU); Arundinaria gigantea (SEMO). Date extremes in Missouri: 1 January–16 December. Discussion. Specimens were reared in the laboratory and immature stages described by Cuda and McPherson (1976). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). In Missouri, this common species feeds on Quercus, Salix, and Ulmus and overwinters under bark (Froeschner 1941).

Chinavia hilaris (Say) Pentatoma hilaris Say, 1831a: Descrip. N. Am. Ins. La., p. 9. Chinavia hilaris: Schwertner and Grazia, 2006, Iher. Ser. Zool. Por. Aleg. 96. Distribution. California and Montana east to Florida and Quebec (Froe- schner 1988). Missouri records are from the City of St. Louis and 59 counties in all six Natural Divisions (MU, SEMO, UMDC). Label data. Biological: Acer negunda, Lespedeza, Salix sp., Solidago, alfalfa, big bluestem, black locust, corn, grain sorghum, impatiens, milkweed, milkweed pod, pin oak leaf, partridge pea, soybeans, sycamore, red clover, flowers and shrubs, 2012 THE GREAT LAKES ENTOMOLOGIST 143 glade edge, grass, with Leptocoris trivittatus, prairie, roadside vegetation, tall grass at forest edge, weeds, blacklight, light trap, lights, malaise trap, mercury vapor light (MU); Arundinaria gigantea (SEMO); in leaf litter under large trees nr river beach (30 Dec.), Urtica, cotton, cavern area, blacklight for SWCB, bollworm pheromone trap, bollworm trap, European corn borer pheromone trap, European corn borer trap, pheromone trap, southwestern corn borer trap, UV trap (UMDC). Date extremes in Missouri: 1 March–30 December (MU, UMDC). Discussion. This species is known commonly as the green stink bug and was described from Missouri specimens. McPherson and Tecic (1997) provided data on the field life history. The biology of this species (asAcrosternum hilare) was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). In Missouri in August, this species was observed mating and feeding on asparagus (Froeschner 1941).

Chinavia pennsylvanica (Gmelin) Cimex viridis pensylvanicus [sic] De Geer, 1773: Mem. Ins. 3: 330, pl. 34, fig 5. Cimex pensylvanicus [sic] Gmelin, 1790: Car. Lin. Syst. Nat. 1: 2148. Chinavia pennsylvanica: Schwertner and Grazia, 2006, Iher. Ser. Zool. Por. Aleg. 96. Distribution. Quebec to Florida west to Iowa and Louisiana (Froeschner 1988). The sole Missouri record is from near Eminence, Shannon County in Ozark Natural Division. Label data. Biological: sweeping weeds. Date collected: 16 June 1940. Discussion. The biology and distribution of this species (as Acrosternum pennsylvanicum) were reviewed by McPherson (1982). This species is rare in Missouri and was collected by sweeping weeds in an overgrown orchard (Froe- schner 1941).

Chlorochroa persimilis Horvath Chlorochroa persimilis Horvath, 1908: An. Mus. Nat. Hung. 6: 555. Distribution. Florida to Quebec west to Arkansas and Alberta (Froe- schner 1988). Missouri records are from the City of St. Louis and six counties in Ozark, Ozark Border, Glaciated Plains, and Osage Plains Natural Divisions. Label data. Biological: red clover (MU); JB [Japanese beetle] trap (MDA); swept from Melilotus (Froeschner 1941). Date extremes in Missouri: 12 April–24 October. Discussion. Several problems exist in the literature concerning species of Chlorochroa from Missouri. Two species, C. sayi (Stål) and C. uhleri (Stål), were included in the species of Pentatomidae known from Missouri (Froeschner 1941); however, each of these was based on only a few old records and identifications prior to the revision by Buxton et al. (1983). All of those Missouri records are now considered to pertain to C. persimilis (D. Thomas, pers. comm.). In addition, Missouri was given as a state in which the primarily western species C. faceta (Say) and C. ligata (Say) occur (Froeschner 1988). These were based on insects collected by Say in his expedition through the Missouri Territory to the Rocky Mountains. Because his notes were stolen, Say had to match specimens to localities from memory. Thus for C. faceta and C. ligata, we consider that either (1) the part of Missouri Territory where these specimens were collected was west of what is now the state of Missouri, (2) they were collected in what is now Missouri, but their ranges have contracted, or (3) they were collected in what is now Missouri but actually were C. persimilis. The latter possibility cannot be confirmed because the types are lost. Further, more recent revisions 144 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 did not list Missouri in the distribution for C. faceta (Thomas 1983) and C. ligata (Buxton et al. 1983). In Missouri, C. persimilis is one of the larger species of Pentatomidae, and was observed on red clover in high abundance (Oetting and Yonke 1971c) and collected on Amsonia spp. (Buxton et al. 1983). This species was reared in the laboratory and life history data were reported by Oetting and Yonke (1971c). The biology and distribution were reviewed by McPherson (1982).

Coenus delius (Say) Pentatoma delia Say, 1831b: Descrip. Het. Hem. N. Am., p. 8. Coenus delius: Stål, 1867, Öfv. K. Svens. Vet.-Akad. Förh. 24(7): 526. Distribution. British Columbia to Quebec south to Texas and Florida (Froeschner 1988). Missouri records are from 19 counties in Ozark, Ozark Border, Osage Plains, and Glaciated Plains Natural Divisions. Label data. Biological: red clover, timothy, white clover, lake vegetation, prairie. Date extremes in Missouri: 9 April–13 November (Oetting and Yonke 1971c, MU). Discussion. The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). The type specimens are from Massachusetts and Missouri. This species is locally scarce and has been swept from Solidago (Froeschner 1941).

Coenus inermis Harris and Johnston Coenus inermis Harris and Johnston, 1936: Ia. St. Coll. J. Sci. 10: 378. Distribution. Kansas, Louisiana (Rider 1996), Oklahoma, Arkansas, and Missouri (Froeschner 1988). Discussion. Froeschner (1941) indicated that C. inermis was collected in Marion County, Arkansas, which is adjacent to Missouri. Therefore, he considered it likely to occur in the south central part of Missouri. Although specific data were not given, Missouri was included in the list of states for this species in the catalog of Heteroptera (Froeschner 1988); however, we have not seen any specimens from Missouri.

Cosmopepla intergressa (Uhler) Eysarcoris melanocephalus: Uhler, 1876, Bull. U.S. Geol. Geogr. Surv. Terr. 1, pl. 9, fig. 7. Cosmopepla humboldtensis Bliven, 1955: Stud Ins. Redwood Emp., 1: 8. Cosmopepla intergressa: McDonald, 1986, J. N. Y. Ent. Soc. 94: 11. Distribution. British Columbia and Montana south to California, Kansas (Froeschner 1941), and Missouri. The sole Missouri record is from Ranken, St. Louis County in the Ozark Border Natural Division. Label data. Date collected: 13 May 1934. Discussion. McDonald (1986) reviewed the genus. This specimen was determined by D. B. Thomas and is the first record of this species from Missouri.

Cosmopepla lintneriana Kirkaldy Cimex carnifex Fabricius, 1798: Ent. Syst., Suppl, p. 535. Cosmopepla lintneriana Kirkaldy, 1909: Cat. Hem. 1: 80. 2012 THE GREAT LAKES ENTOMOLOGIST 145

Distribution. British Columbia to Nova Scotia south to New Mexico, Georgia and Mexico (Froeschner 1988). Missouri records are from the City of St. Louis and 20 counties in Ozark, Ozark Border, Glaciated Plains, Mississippi Lowlands, and Big Rivers Natural Divisions. Label data. Biological: cotton, mullein, red clover, tobacco, flowers, flower garden, flowers & shrubs, grass, vegetation, weeds, wildflowers, blacklight. Date extremes in Missouri: 20 April–20 November. Discussion. McDonald (1986) reviewed the genus. Specimens of C. lintneriana (as C. bimaculata) were reared in the laboratory by McPherson (1976b) and the field life history was studied by McPherson and Tecic (1997). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). Froeschner (1941) reported that this species usually is found by sweeping weeds along streams.

Dendrocoris humeralis (Uhler) Liotropis humeralis Uhler, 1877: Bull. U. S. Geol. Geogr. Surv. Terr. 3: 400. Dendrocoris humeralis: Bergroth, 1891, Rev. Ent. 10: 228. Distribution. Maine to Georgia west to California (Froeschner 1988), and Canada (Paiero et al. 2003). Missouri records are from 11 counties in Ozark, Ozark Border, Glaciated Plains, and Mississippi Lowlands Natural Divisions. Label data. Biological: black locust, black oak, red oak, woods, grass by river, sticky boards in oak hickory canopy, mercury vapor light. Date extremes in Missouri: 2 April–17 October. Discussion. The biology of this species was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). This scarce species in Missouri has been taken on oak trees (Froeschner 1941).

Euschistus ictericus (Linnaeus) Cimex ictericus Linnaeus, 1763: Cent. Ins., p. 16. Euschistus ictericus: Dallas, 1851, List Hem. Brit. Mus. 1: 206. Distribution. Florida, Vermont, and Ontario west to Texas and Colorado (Froeschner 1988). Missouri records are from the City of St. Louis and 5 counties in Ozark, Ozark Border, and Big Rivers Natural Divisions. Label data. Biological: Salix, Japanese beetle trap (Froeschner 1941). Date extremes in Missouri: 18 July–21 October (Froeschner 1941, MU). Discussion. The life history and immature stages of E. ictericus were described by McPherson and Paskewitz (1984b) and mating behavior by Drickamer and McPherson (1992). The biology was reviewed by McPherson and Mohlen- brock (1976) and biology and distribution by McPherson (1982).

Euschistus politus Uhler Euschistus politus Uhler, 1897: Can. Ent. 29: 117. Distribution. Missouri and Arkansas east to New Hampshire and Ten- nessee (Froeschner 1988). Missouri records are from 12 counties in Ozark, Ozark Border, and Big Rivers Natural Divisions. Label data. Biological: Quercus borealis Michr., Quercus stellata Wang, sweeping vegetation, malaise (MU); swept from open woods, Japanese beetle trap (Froeschner 1941). Date extremes in Missouri: 2 May–31 October. 146 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Discussion. Information on laboratory rearing was presented by McPher- son (1974a) and mating behavior by Drickamer and McPherson (1992). The biology of E. politus was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982).

Euschistus servus (Say) Pentatoma serva Say, 1831b: Descrip. Het. Hem. N. Am., p. 4. Euschistus servus: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 26. Distribution. British Columbia and Nova Scotia south to Arizona, Florida, and Mexico (Froeschner 1988). Missouri records are from 84 counties in all six Natural Divisions (MU, UMDC, Froeschner 1941). Label data. Biological: Ambrosia sp., Amorpha canescens, black night- shade Solanum americanum, Daucus carota L. flower, hollyhockAthaea rosea, Lespedeza, Mirabilis nyctaginea, Phlox diverticula, Rudbeckia, Rumex verticil- latus, Salix sp., Solidago spp., alfalfa, alsike clover, bedstraw, Bermuda, cheat, corn, fescue, hop clover, partridge pea, red clover, sunflower, sweet clover, thistle, timothy, tomato plant, vetch roadside, white clover, willow, winter cress, alfalfa/ orchard grass mixture, around pond, emerg. veg., grass, grass/scrub, grasses near a forest, meadow, mixed grass prairie, near shore, prairie, river bottom veg., riverbank, shoreline vegetation, sweep riparian veg., tall grass prairie, tree border at lawn, vegetation, weeds, blacklight, fluorescent light, lights, light trap, malaise, merc. vapor light, USDA Japanese beetle trap (MU); Arundinaria gigantea (SEMO); oak, soybean, sugarbeet, alive with eggs (13 Apr.), bollworm pher. trap, southwestern corn borer trap, Japanese beetle pher. trap (UMDC). Date extremes in Missouri: 31 March–13 November. Discussion. This species is known commonly as the brown stink bug and is represented by two subspecies. The nominate subspecies occurs in the southeastern U. S. and E. servus euschistoides (Vollenhoven) occurs across northern North America, with intergrade forms along a broad band extending from Maryland to Kansas (Sailer 1954). In Illinois, McPherson (1982) determined the entire state south of 40° 30’N (roughly equivalent to the northern border of Missouri) to be populated by intergrade forms, with E. s. euschistoides in the northern part of the state. The distribution of subspecies in Missouri should mirror that of the southern part of Illinois, with most specimens from throughout the state exhibiting combinations of characteristics of both subspecies. Thus, most specimens from Missouri are best referred to as E. s. servus - E. s. euschistoides intergrades. The genus in Mesoamerica was revised by Rolston (1974). The biology of E. servus was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). Information on laboratory life history and behavior were presented by Youther and McPherson (1975), life history, laboratory rearing, and descriptions of immature stages by Munyaneza and McPherson (1994), and mating behavior by Drickamer and McPherson (1992). In Missouri, nymphal stages and adults feed on Verbascum and other plants, and adults overwinter beneath mullein leaves and in grass clumps (Froeschner 1941). This species is very common throughout Missouri.

Euschistus tristigmus (Say) Pentatoma tristigmus Say, 1831b: Descrip. Het. Hem. N. Am., p. 4. Euschistus tristigma: Dallas, 1851, List. Hem. Brit. Mus. 1: 207. Distribution. Nova Scotia and Florida west to Colorado (Froeschner 1988). Missouri records are from the City of St. Louis and 55 counties in all six Natural Divisions (MU, SEMO, UMDC). 2012 THE GREAT LAKES ENTOMOLOGIST 147

Label data. Biological: Commelina diffusa, Fraxinus pennsylvanicus trunk, Quercus velutina, Solidago spp., alfalfa, black-eyed susan, buckeye, corn, fescue, grain sorghum, oak, red clover, red delicious apple tree, smartweed, soybeans, white clover, fallow unweeded field/forest, flowers, grass, herbage, meadow, riparian veg., tree edge, vegetation, weeds, wooded area, blacklight beside pond, blacklight, fluorescent light, light trap, malaise trap, mercury vapor light (MU); Arundinaria gigantea (SEMO); pheromone trap (UMDC). Date extremes in Missouri: 28 February–26 October. Discussion. This species is known commonly as the dusky stink bug and is represented by two subspecies. McPherson (1982) suggested that populations north of 41o N were E. t. luridus Dallas and those to the south were the nominate subspecies. He also provided a taxonomic key to distinguish them from each other and from congeners. The northern border of Missouri is at latitude 40° 35’, thus, based on this geographic delineation, all specimens from Missouri are south of 41o latitude and should be the nominate subspecies. However, specimens from areas near the line, including the northern counties of Missouri, should be examined closely to determine to which subspecies they belong. The genus in Mesoamerica was revised by Rolston (1974). The life history of E. t. tristigmus based on laboratory rearing and field observations were presented by McPherson (1975b) and mating behavior by Drickamer and McPherson (1992). The biology of E. tristigmus was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). McPherson (1974b, 1975a, 1976a 1979c,e) and McPherson and Paskewitz (1982) conducted a series of controlled laboratory experiments on the effects of photoperiod on adult morphology. Adults of this species have been found feeding Ambrosia and adults and nymphs on Quercus and blackberries (Froe- schner 1941).

Euschistus variolarius (Palisot) Pentatoma variolaria Palisot de Beauvois, 1817: Ins. Rec. Afr. Am., p. 149. Euschitus [sic] punctipes: Rathvon, 1869, Hist. Lanc. Co. Pa., p. 549. Euschistus variolarius: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 26. Distribution. Quebec and Florida west to British Columbia and Utah (Froeschner 1988). Missouri records are from the City of St. Louis and 93 counties in all six Natural Divisions. Label data. Biological: Aesculus, Aster, Baptisia alba, Baptisia leu- cantha, Bidens at pond margin, Lonicera japonica, Matricaria sp, Monarda futulosa, Rudbeckia, Salix flowers, Shrankia, Solidago, Verbascum thaspi, Verbascum, Zinnia, adino clover, alfalfa, alsike, barley, cheat, corn, fescue, hickory tree, hop clover, milkweed, partridge pea, peaches, red clover, soybeans, timothy, wheat, white clover, winter cress, around pond, building wall (October), field with many types of weeds, glade, grass field, grass, ground vegetation, lake vegetation, meadow, native grass plot, prairie meadow, prairie, roadside vegetation, tree border at lawn, under bark of dead tree (Nov. 10), weeds, yellow flower, blacklight, light trap, lights, malaise trap, mercury vapor lt. (MU); tobacco budworm trap (UMDC). Date extremes in Missouri: 26 February–1 December. Discussion. Comparative information on mating behavior was presented by Drickamer and McPherson (1992). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). This very common and widespread species overwinters as adults under leaves and in grass clumps (Froeschner 1941). 148 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Halyomorpha halys (Stål) Pentatoma halys Stål, 1855: Öfv. K. Svens. Vet.-Akad. Förh. 12(4): 182. Halyomorpha picus: auctorum (nec Fabricius, 1794: 115). Halyomorpha halys: Josifov and Kerzher, 1978, Frag. Faun. 23: 172. Distribution. Maine to Florida west to Illinois (Zhu et al. 2012) and Mississippi, with a disjunct population in California, Oregon, and Washington. The sole Missouri record to date is from Maryland Heights, St. Louis County in the Ozark Border Natural Division. Label data. Date of collection: 16 March 2012. Discussion. Halyomorpha halys is known commonly as the brown marmorated stink bug. This invasive species is native to eastern Asia where it is known from approximately 300 host plant species and is a pest on tree fruits and soybeans (Nielsen and Hamilton 2009, and citations therein). It also is a nuisance pest as it enters homes in large numbers at the onset of cold weather (Hoebeke and Carter 2003). This species was introduced recently to Europe (Wermelinger et al. 2008) and New Zealand (Harris 2010). It was first found in the U. S. in Pennsylvania in 1996 (Hoebeke and Carter 2003) and its range in North America has been expanding rapidly. Nielsen and Hamilton (2009) reported on the life history of H. halys in the northeastern United States. The entire state of Missouri is in the area considered to be highly suitable for invasion by this species (Zhu et al. 2012). The first live specimen in Missouri, a female, was collected recently on a curb near an interstate highway near St. Louis. This is the first record of this species from Missouri.

Holcostethus limbolarius (Stål) Peribalus limbolarius Stål, 1872: K. Svens. Vet.-Akad. Handl. 10(4): 34. Holcostethus limbolarius: Kirkaldy, 1909, Cat. Hem. 1: 48. Distribution. Maine to Georgia west to British Columbia and Arizona (Froeschner 1988). Missouri records are from the City of St. Louis and 49 counties in all six Natural Divisions (MU, UMDC). Label data. Biological: Cleoma serrulata, Euthamia gymnospermoides, Solidago & Eupatorium, alfalfa, bean, corn & bean, goldenrod, meadow goldenrod, meadow-smooth aster, red clover, wheat, white clover, brush & flower, fence row, garden turnip broccoli etc, grass, leaf litter, oak-hickory 2nd growth, prairie, red clover/fescue, blacklight, lights, malaise (MU); sugar beet (UMDC). Date extremes in Missouri: 14 January–7 December. Discussion. The genus in North America was revised by McDonald (1974). Biological information was provided by Oetting and Yonke (1971c). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). Froeschner (1941) considered this to be the second most common pentatomid species in the state. He reported that its host plants include Solidago and Polygonum and that it overwinters in protected places, such as under mullein leaves and in grass clumps.

Hymenarcys nervosa (Say) Pentatoma nervosa Say, 1831b: Descrip. Het. Hem. N. Am., p. 9. Hymenarcys perpunctata Amyot and Serville, 1843: Hist. Nat. Ins. Hem., p. 124. Hymenarcys nervosa: Walker, 1867, Cat. Hem. Het. Brit. Mus. 2: 283. Distribution. Quebec to Florida west to Iowa and Texas (Froeschner 1988). Missouri records are from the City of St. Louis and 52 counties in all six Natural Divisions (MU, UMDC). 2012 THE GREAT LAKES ENTOMOLOGIST 149

Label data. Biological: Lespedeza, Verbascum, alfalfa, blue grass, goldenrod, hop clover, low hop clover, mullein leaves, red clover, sweet clover, white clover, under bark (14 Apr.), under log (9 Apr.), vegetation, woods, Berlese sample, malaise, mercury vapor light (MU); weed (UMDC). Date extremes in Missouri: 7 January–3 December. Discussion. The genus was reviewed by Rolston (1973). The biology of H. nervosa was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). In Missouri, this species overwinters as adults in grass clumps and under rocks and mullein leaves (Froeschner 1941).

Mcphersonarcys aequalis (Say) Pentatoma aequalis Say, 1831b: Descrip. Het. Hem. N. Am., p. 7. Hymenarcys aequalis: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 30. Mcphersonarcys aequalis: Thomas, 2012, Gr. Lakes Ent. 45: 127. Distribution. Massachusetts to Montana south to Florida, Texas, and Mexico (Froeschner 1988). Missouri records are from 12 counties in Ozark, Glaciated Plains, and Mississippi Lowlands Natural Divisions. Label data. Biological: red clover, weeds, under bark (16 Nov.) (MU); under bark of standing tree (Feb.), weedy fields (Froeschner 1941). Date extremes in Missouri: 1 January-16 November. Discussion. The genus (as Hymenarcys) was reviewed by Rolston (1973). Descriptions of immature stages and biology were given by Oetting and Yonke (1971b). The biology of this species was reviewed by McPherson and Mohlen- brock (1976) and biology and distribution by McPherson (1982).

Mecidea major Sailer Mecidea longula: Uhler, 1876, Bull. U.S. Geol. Geogr. Surv. Terr. 1: 283 (in part). Mecidea major Sailer, 1952: Proc. U.S. Nat. Mus. 102: 478, 486. Distribution. Most states in a line from Arizona to Illinois (Froeschner 1988). Missouri records are from 3 counties in Ozark, Ozark Border, and Osage Plains Natural Divisions. Label data. Biological: grass field. Date extremes in Missouri: 10 Sep- tember–10 October. Discussion. Sailer (1952) reported this species from Missouri. Com- parative laboratory rearing was presented by Bundy et al. (2005). Biology and distribution were reviewed by McPherson (1982).

Mecidea minor Ruckes Mecidea longula: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 17 (in part). Mecidea minor Ruckes, 1946a: Bull. Brook. Ent. Soc. 41: 87. Distribution. South Dakota and Iowa southwest to California, Texas, and Mexico (Froeschner 1988). Discussion. Sailer (1952) reported this species from Missouri. Compara- tive laboratory rearing was presented by Bundy et al. (2005), and life history and descriptions of immature stages by Bundy and McPherson (2011). Biology and distribution were reviewed by McPherson (1982). We have not seen specimens from Missouri and its inclusion here is based on Sailer (1952). 150 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Menecles insertus (Say) Pentatoma inserta Say, 1831b: Descrip. Het. Hem. N. Am., p. 6. Menecles insertus: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 31. Distribution. Quebec to New Jersey west to California (Froeschner 1988). Missouri records are from 10 counties in Osage Plains, Ozark Border, and Big Rivers Natural Divisions (MU, UMDC). Label data. Biological: sandy bank, light, light trap (MU); grass (UMDC). Date extremes in Missouri: 4 April–11 November. Discussion. Menecles insertus was described from specimens from Arkan- sas and Missouri. The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). Froeschner (1941) reported this species usually is found in association with various oaks.

Mormidea lugens (Fabricius) Cimex lugens Fabricius, 1775: Syst. Ent., p. 716. Mormidea lugens: Walker, 1868, Cat. Hem. Brit. Mus. 3: 551. Distribution. Nova Scotia to Manitoba south to Florida, Texas and Mexico (Froeschner 1988). Missouri records are from the City of St. Louis and 59 counties in Ozark, Ozark Border, Osage Plains, Glaciated Plains, and Mis- sissippi Lowlands Natural Divisions. Label data. Biological: alfalfa, fescue, hop clover, oak foliage, partridge pea, red clover, tall fescue, wheat, white clover, meadow, near pond, night sweeping native prairie, prairie, sweeping grasses, vegetation, weeds, malaise trap (MU); sugar beet (UMDC). Date extremes in Missouri: 28 January–10 December. Discussion. Information on biology was provided by Oetting and Yonke (1971c) and McPherson (1974a). The biology of this species was reviewed by McPherson and Mohlenbrock (1976). Nymphs have been found in Missouri from June through August (Froeschner 1941).

Murgantia histrionica (Hahn) Strachia histrionica Hahn, 1834: Wanz. Ins. 2: 116. Murgantia histrionica: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 37. Distribution. Connecticut to California south to Florida and Mexico (Froeschner 1988). Missouri records are from 21 counties in all six Natural Divisions (MU, UMDC). Label data. Biological: beans, cabbage, cleome, kale leaf, squash, turnips (MU); collards, mustard, sweeping weeds (UMDC). Date extremes in Missouri: 4 April–1 November (Froeschner 1941, MU). Discussion. This species is known commonly as the harlequin bug and can be a serious pest of cabbage (Froeschner 1941). The biology and distribution were reviewed by McPherson (1982).

Neottiglossa cavifrons Stål Neottiglossa cavifrons Stål, 1872: K. Svens. Vet.-Akad. Handl. 10(4): 18. Distribution. Oregon to Arizona east to Virginia and Georgia (Froeschner 1988). Missouri records are from the City of St. Louis and 13 counties in Ozark, Ozark Border, Glaciated Plains, and Mississippi Lowlands Natural Divisions. 2012 THE GREAT LAKES ENTOMOLOGIST 151

Label data. Biological: white clover, prairie. Date extremes in Missouri: 6 May–27 October. Discussion. This genus in North America was reviewed by Rider (1990). This species was reported from Missouri by Froeschner (1941) and Oetting and Yonke (1971c). Descriptions of immature stages and biological information was provided by Oetting and Yonke (1971c). The biology was reviewed by McPher- son and Mohlenbrock (1976) and biology and distribution by McPherson (1982). This species often is found with its congener, N. sulcifrons, but is slightly more numerous (Froeschner 1941).

Neottiglossa sulcifrons Stål Neottiglossa sulcifrons Stål, 1872: K. Svens. Vet.-Akad. Handl. 10(4): 18. Distribution. New Jersey to Georgia west to Utah and New Mexico (Froeschner 1988), and Canada (Scudder 2004). Missouri records are from 23 counties in all six Natural Divisions. Label data. Biological: Andropogon, prairie, sandbar, malaise. Date extremes in Missouri: 1 May–1 December. Discussion. This genus in North America was reviewed by Rider (1990). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). This common species has been collected in Missouri from dry, weedy fields and open woods (Froeschner 1941).

Neottiglossa undata (Say) Pentatoma undata Say, 1831b: Descrip. Het. Hem. N. Am., p. 8. Neottiglossa undata: Uhler, 1871, Proc. Bost. Soc. Nat. Hist. 14: 395 (in part). Distribution. British Columbia to Quebec south to California, Nebraska (Froeschner 1988) and North Carolina (Torre Bueno 1913). The sole Missouri record is from Columbia, Boone County in the Ozark Border Natural Division. Label data. Biological: sweep. Date collected: 28 August 1989. Discussion. The biology and distribution were reviewed by McPherson (1982). This specimen was determined by D. Rider as part of his 1990 review of North American species, and is the first record of this species from Missouri.

Oebalus pugnax (Fabricius) Cimex pugnax Fabricius, 1775: Syst. Ent., p. 704. Oebalus typhoeus: Stål, 1868, K. Svens. Vet.-Akad. Handl. 7(11): 27. Oebalus pugnax: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 22. Distribution. Connecticut to Florida west to Minnesota, Arizona, and Mexico (Froeschner 1988). Missouri records are from the City of St. Louis and 47 counties in Ozark, Ozark Border, Osage Plains, Glaciated Plains, and Mis- sissippi Lowlands Natural Divisions. Label data. Biological: Passiflora, alfalfa, barley, hop clover, partridge pea, ragweed, red clover, rice, sorghum, timothy, white clover, bush & flower, grass, grass/shrub, meadow, prairie, roadside ditch, soybeans & corn, tall grass mating (1 Sept.), vegetation, weeds, blacklight, fluorescent light, light trap, malaise trap (MU); Arundinaria gigantea (SEMO); sugar beet, field margin, black cutworm trap, bollworm trap, bollworm pheromone trap, European corn borer trap, light trap, southwestern corn borer trap, soybean looper trap (UMDC). Date extremes in Missouri: 15 May–28 October (UMDC, SEMO). 152 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Discussion. This species is known commonly as the rice stink bug and can reach population densities sufficiently high to cause economic damage to rice by reducing quality or yield. The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). Froe- schner (1941) reported that this species (as Solubea pugnax) tends to be more common in the southern counties of Missouri, that it was observed feeding on the grass Setaria, and that adults overwinter in grass clumps.

Parabrochymena arborea (Say) Pentatoma arborea Say, 1825: J. Acad. Nat. Sci. Phila. 4: 311. Parabrochymena arborea: Larivière, 1992, Mem. Ent. Soc. Can. 163: 7. Distribution. Quebec to Florida west to Kansas, New Mexico, and Mexico (Froeschner 1988). Missouri records are from the City of St. Louis and 22 counties in Ozark, Ozark Border, and Glaciated Plains Natural Divisions (MU, SEMO, UMDC). Label data. Biological: Quercus imbricaria Mich., goldenrod, oak, pin oak, red delicious apple, picnic table, sandy bank, sweeping grass (MU); hybrid elm (UMDC). Date extremes in Missouri: 12 April–5 November (UMDC, SEMO). Discussion. This species was described from Missouri (Say 1825). The biology (as Brochymena arborea) was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). This species was considered to be locally uncommon in Missouri by Froeschner (1941).

Piezodorus guildinii (Westwood) Raphigaster guildinii Westwood, 1837: Hope Cat., p. 31. Piezodorus guildingi [sic]: Uhler, 1894, Proc. Zool. Soc. Lond., p. 175. Distribution. Argentina (Panizzi and Slansky 1985) north to New Mexico, Georgia (Froeschner 1988), and Missouri (Tindall and Fothergill 2011). Mis- souri records are from 2 counties in the Mississippi Lowlands Natural Division (Tindall and Fothergill 2011). Label data. Biological: black light. Date extremes in Missouri: 10–11 October. Discussion. This predominantly Neotropical species is known commonly as the redbanded stink bug and a pest on soybeans. Its biology and ecology on soybeans were studied by Arroyo and Kawamura (2003) and host plant information was provided by Panizzi and Slansky (1985). More recent accounts of this species on soybeans in the U. S. have been reported in Arkansas (Smith et al. 2009), Louisiana (Temple 2011), and Missouri (Tindall and Fothergill 2011).

Prionosoma podopioides Uhler Prionosoma podopioides Uhler, 1863: Proc. Ent. Soc. Phila. 2: 364. Distribution. Arizona to British Columbia east to Michigan and Illinois (Froeschner 1988). Missouri records are from 7 counties in Ozark, Ozark Border, Glaciated Plains, and Osage Plains Natural Divisions. Label data. Date extremes in Missouri: 11 February–27 November (MU), and to December (Froeschner 1941). Discussion. McPherson and Cuda (1975) caught an adult female and kept it alive in a mason jar for 1.5 months by feeding it Plantago aristata Michaux. The biology and distribution of this species were reviewed by McPherson (1982) and Wheeler (1988). Froeschner (1941) reported Missouri records of this scarce 2012 THE GREAT LAKES ENTOMOLOGIST 153 species from fields and open woods whereP. aristata was common, and that an adult was collected beneath bark in December.

Proxys punctulatus (Palisot) Halys punctulatus Palisot de Beauvois, 1817: Ins. Rec. Afr. Am., p. 188. Proxys punctulatus: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 29. Distribution. Pennsylvania and Missouri south to Florida, Texas (Froe- schner 1988), and Brazil (Gomez and Mizell 2012). Missouri records are from 13 counties in Ozark, Ozark Border, Osage Plains, and Mississippi Lowlands Natural Divisions (MU, SEMO, UMDC). Label data. Biological: blacklight, lights (MU); soybeans, soybean looper trap, UV trap (UMDC); Japanese beetle trap (Froeschner 1941). Date extremes in Missouri: 23 May–29 September. Discussion. This species is known commonly as the black stink bug and was reported from Japanese beetle traps in St. Louis by Froeschner (1941). The life history and laboratory rearing of this species were reported by Vangieson and McPherson (1975). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982).

Tepa vanduzeei Rider Tepa vanduzeei Rider, 1986: J. N. Y. Ent. Soc. 94: 555. Distribution. California to South Dakota south to Mexico (Froeschner 1988). The sole Missouri record is from Metz, Vernon County in the Osage Plains Natural Division. Label data. Date collected in Missouri: 21 April 1940. Discussion. The only known Missouri specimen is a paratype. This specimen originally was reported as Thyanta rugulosa (Say) (now Tepa rugulosa) by Froeschner (1941), and was revised by Rider (1986).

Thyanta calceata (Say) Pentatoma calceata Say, 1831b: Descrip. Het. Hem. N. Am., p. 8. Thyanta calceata: Barber, 1911, J. N. Y. Ent. Soc. 19: 108. Distribution. Virginia to Florida west to Texas (Froeschner 1988). Mis- souri records are from the City of St. Louis and 41 counties in all six Natural Divisions (MU, MDA, Rider and Chapin 1992). Label data. Biological: Commelina diffusa, Rhus aromaticus, alfalfa, hop clover, low hop clover, red clover, timothy, flowers, grass, herbage, prairie, soybeans & corn, vegetation, light trap, malaise, mercury vapor light. Date extremes in Missouri: 4 April–7 November. Discussion. The biology of this species was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). McPherson (1977a, 1978), and McPherson et al. (1983) conducted controlled laboratory experiments on the effects of photoperiod on adult morphology. The immature stages were described by Paskewitz and McPherson (1982) and data on seasonal dimorphism was presented by McPherson (1977b). Froeschner (1941) considered this species to be scarce in Missouri and that most specimens were collected from Ozark woods, although that was based on records from only 6 counties. The species appears to be widespread throughout the state based on records from current museum holdings. 154 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Thyanta custator accerra McAtee Cimex custator Fabricius, 1803: Syst. Rhyn. p. 164. Thyanta custator: Stål, 1860, K. Svens. Vet.-Akad. Handl. 2(7): 58. Thyanta custator var. accerra McAtee, 1919: Bull. Brook. Ent. Soc. 14: 16. Distribution. Missouri records are from the City of St. Louis and 88 counties in all six Natural Divisions (MU, SEMO, UMDC). Label data. Biological: Acacia greggii, Chrysanthemum leucanthemum L. flower, feeding on fruit of Cirsium vulgare (Shari) tenore, Cleoma serrulata, Commelina diffusa, Gallardia aristata, Lespedeza, Medicago sativa L., Mirabilis nyctaginea, Rhus aromaticus, Rudbeckia, Thuja, alfalfa, aster, blue grass, buck- brush, goldenrod, hop clover, low hop clover, marigold, partridge pea, peaches, red clover, soybeans, sweet clover, timothy, white clover, flowers, herbage, pasture, prairie, soybean & corn, sweeping flowers, sweeping grasses, vegetation, blacklight, incandescent light, light trap, lights, malaise, mercury vapor light (MU); mustard, soybeans with small pods, wheat, mixed sample from soybean and cotton, black cutworm [trap], blacklight for southwestern corn borer, bollworm trap, bollworm pher. trap (UMDC). Date extremes in Missouri: 4 April–11 December (MU, SEMO). Discussion. This subspecies is known commonly as the redshouldered stink bug (McPherson 1982). There has been confusion regarding the taxonomy of T. custator and the status of what is now regarded as the subspecies T. custator accerra McAtee. The latter has been variously recognized as a variety of T. custator, a full species, and a subspecies of T. pallidovirens (Rider and Chapin 1992). The range of the nominate subspecies is along the Gulf and Atlantic coastlines and does not extend into Missouri. Thus, all specimens of T. custator from Missouri are T. c. accerra. The biology (as T. pallido-virens accerra) was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution (as T. accerra) by McPherson (1982). Comparative information on mating behavior was presented by Dricka- mer and McPherson (1992). McPherson (1979d) conducted controlled laboratory experiments on the effects of photoperiod on adult morphology. Oetting and Yonke (1971c) provided supplemental rearing data based on Missouri specimens, reported parasitoids of this species in Missouri, and observed feeding on mullein, soybeans, and red clover. Oetting and Yonke (1971c) further reported that although T. custator accerra attacks some crops, it density does not reach economic injury levels. Froeschner (1941) considered this to be the most common stink bug in Missouri and reported that adults overwinter in grass clumps and under mullein leaves.

Trichopepla semivittata (Say) Pentatoma semivittata Say, 1831b: Descrip. Het. Hem. N. Am., p. 9. Trichopepla semivittata: Uhler, 1871, Proc. Bost. Soc. Nat. Hist. 14: 96. Distribution. Quebec to Florida west to Colorado, Texas, and Mexico (Froeschner 1988). Missouri records are from the City of St. Louis and 30 counties in Ozark, Ozark Border, Osage Plains, Glaciated Plains, and Mississippi Lowlands Natural Divisions (MU, MDA). Label data. Biological: Eupatorium & asters, alsike, alfalfa, meadow goldenrod, Queen Anne’s lace, red clover, white aster, wild carrot, low woodland veg., garden turnip brussel sprouts, prairie, weeds (MU); JB [Japanese beetle] trap, USDA traps (MDA). Date extremes in Missouri: 27 April–26 October (MU), and to late November (Froeschner 1941). Discussion. The genus in North America was revised by McDonald (1976). T. semivittata was reared in the laboratory on wild carrot (McPherson 1972). 2012 THE GREAT LAKES ENTOMOLOGIST 155 The biology of this species was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). In Missouri, this species tends to be found only within 18 inches from the ground (Froeschner 1941).

SUBFAMILY – PODOPINAE Amaurochrous brevitylus Barber and Sailer Amaurochrous brevitylus Barber and Sailer, 1953: J. Wash. Acad. Sci. 43: 160. Distribution. Massachusetts and Quebec south to Pennsylvania and west to Nebraska, Kansas, and Arizona (Froeschner 1988). Missouri records are from 7 counties in Glaciated Plains, Ozark Border, and Mississippi Lowland Natural Divisions (MU, SEMO). Label data. Biological: prairie, under debris, light trap, malaise, sweep, vacuum sweep (MU); Arundinaria gigantea (SEMO). Date extremes in Missouri: 28 January–23 December. Discussion. The turtle bugs were revised by Barber and Sailer (1953). The biology and distribution of this mostly northern species was reviewed by McPherson (1982). Although in Illinois A. brevitylus is known from only the northern half of the state, the few Missouri records are from throughout the state, including the bootheel. This is the first published account of this species from Missouri.

Amaurochrous cinctipes (Say) Tetyra cinctipes Say, 1828: Phila. Mus. 3: plate 43. Podops (Amaurochrous) cinctipes: Stål, 1872, K. Svens. Vet.-Akad. Handl. 10(4): 15. Amaurochrous cinctipes: Schouteden, 1905, Gen. Ins. 30: 33. Distribution. Quebec and South Carolina west to Nebraska and Texas (Froeschner 1988). Missouri records are from the City of St. Louis and 6 counties in the Glaciated Plains, Osage Plains, Ozark Border, and Mississippi Lowlands Natural Divisions. Label data. Biological: hollow weeds, prairie, USDA traps (MU); Japanese beetle traps (Froeschner 1941). Date extremes in Missouri: 6 May–16 October. Discussion. The turtle bugs were revised by Barber and Sailer (1953). This species was reared in the laboratory and immature stages were described by McPherson and Paskewitz (1984a). The biology was reviewed by McPherson and Mohlenbrock (1976) and biology and distribution by McPherson (1982). Froeschner (1941) reported this species occurs in low marshy areas or among weeds bordering bodies of water and that it is scarce in Missouri.

Species Of Possible Occurrence In Missouri Chlorochroa belfragei Stål. This apparently northern species is considered rare and has been recorded from Canada, Illinois, Iowa, and Nebraska (McPher- son 1982). Nothing is known about the life history of C. belfragei. In Missouri, its occurrence is most probable in the Glaciated Plains Natural Division. Nezara viridula (Linnaeus). The southern green stink bug is a major pest of soybeans in the southeastern U. S. and is known from as far north as Pine Hills, Union County, Illinois (McPherson and Cuda 1974) and Oklahoma (Anonymous 1974). Its occurrence in the southern part of Missouri, especially in the Mississippi Lowlands Natural Division, is probable. 156 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Acknowledgments Foremost, we thank J. E. McPherson for his encouragement, insight, spirit, and friendship over many years. It is solely because of his influence on us that this paper was published. We thank Collin Wamsley and Sarah Phipps, Missouri De- partment of Agriculture, for access to their pentatomid collection and for donating the state record specimen of Halyomorpha halys to the Enns Entomology Museum. We also thank Kelly Tindall, University of Missouri Delta Center - Portageville, for providing records of pentatomids from the Delta Center collection, and Lisa Miehls for data collection from MU specimens. We are grateful to Richard Thom, Missouri Department of Conservation, and James Wilson, University of Missouri - St. Louis, for permission to reprint the Missouri Natural Divisions figure. We thank Donald B. Thomas, USDA - Kerrville, TX, and David A. Rider, University of North Dakota, for helpful comments during preparation of this manuscript.

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Abstract The exoskeletal morphology of the structures associated with the production of substrate vibrations used for communication was examined for members of three guilds of sap-feeding insects on the cordgrasses Spartina alterniflora Loisel, S. patens (Aiton) Muhl., and S. pectinata Link (Poaceae: Chloridoidea). Measurements of the second abdominal sternite and its apodemes, the “tymbal,” were made for the males of 14 species of planthoppers and 2 species of leafhoppers (Hemiptera: Auchenorrhyncha: Caliscelidae, Cicadellidae, Delphacidae, Derbidae). Morphometric comparisons revealed significant differences among the insect species on each of the cordgrass species. If tymbal morphology reflects definitive features of the vibrational signals then coexistence by the members of each sap-feeding guild is likely fostered by partitioning the “substrate resource.” Tymbal morphology may be a valuable tool for determining the presence of sibling species and for providing insights regarding behavior, ecology, and evolution of these insects.

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The sap-feeding insects of three species in the grass genus Spartina (Poaceae: Chloridoidea), S. alterniflora Loisel, S. patens (Aiton) Muhl., and S. pectinata Link, have been the focus of extensive ecological and behavioral studies (e.g., Heady and Wilson 1990, Holder 1990, Heady and Denno 1991, Holder and Wilson 1992, Denno and Peterson 2000). Each of these Spartina species harbors a guild of monophagous or oligophagous sap-feeding planthoppers and leafhoppers (Hemiptera: Auchenorrhyncha) – four species on S. alterniflora, ten species on S. patens, and five species on S. pectinata (Table 1). Species of Spartina also harbor other sap-feeding hemipterans such as aphids and scale insects (Sternorrhyncha: Aphididae, Coccoidea), and seed bugs (Ischnodemus spp., Blissidae) and stink bugs (Rhytidolomia saucia (Say), Pentatomidae) (Johnson and Knapp 1996). As well, there are generalist leafhoppers (Cicadellidae) that occasionally feed on these grasses (Whitcomb et al. 1987). Coexistence of seven of the sap-feeding insects on S. patens apparently results from differences among the species in microhabitat occupancy, phenology, and body size (Denno 1980). Substrate-borne vibrational signals can serve several functions including species recognition and mate location, recruitment to feeding locations, and predator avoidance (Cocroft and McNett 2006, Virant-Doberlet et al. 2006, Hill 2009). These signals are employed by numerous leafhopper, treehopper, and planthopper species (Cocroft 1999, Drosopolous and Claridge 2006, Zezlina and Virant-Doberlet 2007), and a cicada species (Claridge et al. 1999). During courtship, signals are produced by males then detected by females, which typically respond with a simpler signal (Ichikawa and Ishii 1974, Ichikawa 1976). The signals aid in locating potential mates, are species-specific, and serve to maintain prezygotic reproductive isolation (Inoue 1982, Heady and Denno 1991,

1Department of Biology, University of Central Missouri, Warrensburg, MO 64093 (e-mail: [email protected]). 2012 THE GREAT LAKES ENTOMOLOGIST 165

Table 1. The sap-feeding Auchenorrhyncha guilds of three species of Spartina (Denno 1977, 1980; Holder 1990).

Taxon S. alterniflora S. patens S. pectinata

Caliscelidae Aphelonema - simplex Uhler simplex Uhler

Delphacidae Delphacodes penedetecta Beamer detecta (Van Duzee) -

Megamelus n. sp. lobatus Beamer metzaria Crawford

Neomegamelanus - spartini (Osborn) - elongatus (Ball)

Prokelisia dolus Wilson - crocea (Van Duzee) marginata (Van Duzee)

Tumidagena - minuta McDermott - terminalis (Metcalf)

Derbidae Anotia - - fitchi (Van Duzee)

Cicadellidae Amplicephalus - simplex (Van Duzee) -

Destria - bisignata (Sanders & DeLong) -

Neohecalus - lineatus (Uhler) lineatus (Uhler)

Gillham and de Vrijer 1995, Langellotto and Denno 2001). Reproductive isolation has been documented for sibling species of Enchenopa, Nilaparvata, and Ribautodelphax all of which are monophagous on different host plants (Claridge 1985a,b; de Winter and Rollenhagen 1990; Claridge and deVrijer 1994; Hunt 1994; Rodriguez et al. 2004) or sibling species of Prokelisia which occur on the same host species (Heady and Denno 1991). The vibrational signals are produced by rapid movements of the entire abdomen or by movement of internal abdominal exoskeletal structures col- lectively referred to as tymbals (Ossiannilsson 1949, Mitomi et al. 1984). The structures responsible for producing signals have not been determined in some taxa (e.g., flatid planthoppers). In delphacid planthoppers, signals are produced by tymbals (“Singapparat” or “drumming organs” of Asche (1985) and Ossian- nilsson 1949) which consist of exoskeletal modifications of the metapostnotum and the tergites and sternites of the first and second abdominal segments. The morphology of these exoskeletal tymbal elements and their associated muscles have been described and illustrated for Dicranotropis hamata (Boheman) and briefly described for 25 additional species of delphacids, one cixiid, and one issid by Ossiannilsson (1949). Detailed descriptions and illustrations of the exoskeletal morphology and anatomy of the muscles of Nilaparvata lugens (Stål) were made by Mitomi et al. (1984). The exoskeletal morphology and mechanism for 166 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 sound production in N. lugens was described by Zhang et al. (1988). The tymbals of 36 species of delphacids were illustrated by Asche (1985) who used major structural differences in the tymbals in his phylogenetic analysis of the family. Differences in the vibrational signals likely result from differences in tymbal exoskeleton morphology, muscular morphology, and/or neuromuscular physiology. Tymbal morphology differed significantly in four out of five species of Prokelisia with the greatest differences in tymbal structures between Prokeli- sia marginata (Van Duzee) and Prokelisia dolus Wilson, which share the same host plant, Spartina alterniflora (Miller and Wilson, 1999) (Table 1). These two species of Prokelisia produce species specific vibrational signals that serve to prevent hybridization (Heady and Denno 1991). Several species of sap-feeders that each produce vibrational signals critical for reproduction occupy the same Spartina host plants (Table 1). Interference in courtship behavior would be possible among species producing vibrational signals at the same time unless there are significant differences in those signals. As tymbal morphology and signal production are possibly linked, I compared the morphology of species occurring on each of the three Spartina species to determine if there are significant structural differences among them.

Materials and Methods The exoskeletal morphology of the tymbals of 14 species of planthoppers and 2 species of leafhoppers was examined. Specimens were obtained from collections at the University of Central Missouri and the University of Maryland. Specimens were collected in the following localities: Delphacodes penedetecta Beamer, D. detecta (Van Duzee), Megamelus n. sp., M. lobatus Beamer, Neome- gamelanus elongatus (Ball), N. spartini (Osborn), Prokelisia dolus, P. marginata, Tumidagena minuta McDermott, T. terminalis (Metcalf), and Amplicephalus simplex (Uhler) in Ocean Co., New Jersey; Neohecalus lineatus (Uhler) in Ocean Co., New Jersey and Pettis Co., Missouri; Anotia fitchi(Van Duzee) in Johnson Co., Missouri; Megamelus metzaria Crawford and P. crocea (Van Duzee, in Johnson Co. and Pettis Co., Missouri; Aphelonema simplex Uhler in Pettis Co, Missouri, and Hancock Co. and Tuscola Co., Michigan; P. dolus in Piaquemines Parish, Louisiana and Dauphin Island, Alabama. The abdomens of four males and one female of each species were removed and cleared in 10% potassium hydroxide for ca. 24 h, washed for 10 min in water, then placed in glycerol for examination. If tymbal morphology was difficult to discern after clearing, the specimen was stained with lignin. After examination, each abdomen was placed in a glycerol filled genitalia vial and pinned under the specimen from which it came. There were no obvious differences among males of the different species in the morphology of the tymbal elements of the metapostnotum and first abdominal sternite. The apodemes of the second abdominal sternite were dissimilar and were the focus of description, illustration, and morphometric analyses. Females had no discernible development of any structures thus were not described, illustrated, nor measured. Specimens were oriented so that the apodemes of the second abdominal sternite were visible in caudal view. Measurements of the apodemes were made using an ocular micrometer and included: 1) distance between the apodeme bases, 2) length of the apodemes from base to apex, 3) distance between apices of the apodemes, 4) width of the abdominal sternite, 5) depth of the middle of the abdominal sternite, and 6) depth of the abdominal sternite under the apodemes (see Miller and Wilson 1999). Morphometric differences were analyzed using one way analysis of variance (ANOVA) for each species inhabiting the same host plant. For those morphometric features found to be significantly different, pairwise comparisons of species were conducted using Tukey’s Honestly Significant Difference test (Roscoe 1975). 2012 THE GREAT LAKES ENTOMOLOGIST 167

Results Descriptions of the tymbals of the sap-feeding guild of Spartina alterniflora. Delphacodes penedetecta (Fig. 1A, Table 2). The tymbal apodemes were slender, elongate, darker than the abdominal sternum, rounded at the apices and converged toward the apices. The abdominal sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was short and sinuate on the dorsal aspect and rounded laterally. Megamelus n. sp. (Fig. 1B, Table 2). The tymbal apodemes were slender, elongate, darker than the abdominal sternum, rounded at the apices, and subparallel to each other. The abdominal sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was sinuate on the dorsal aspect and rounded laterally. Prokelisia dolus (Fig. 1C, Table 2). The tymbal apodemes were very slender, elongate, darker than the abdominal sternum, rounded at the apices, and diverged from the bases to the apices. The abdominal sternum narrowed between the apodemes and the portion extending lateral from the base of each apodeme was sinuate on the dorsal aspect and tapered laterally. Prokelisia marginata (Fig. 1D, Table 2). The tymbal apodemes were broad and subtriangular and rounded at the apices. The position of the apodemes did not diverge towards the apices. The abdominal sternum was extremely narrow between the apodemes and the portion extending laterally from the base of the apodemes was relatively short, concave on the dorsal aspect and rounded laterally. Descriptions of the tymbals of the sap-feeding guild of Spartina patens. Delphacodes detecta (Fig. 2B, Table 3). The tymbal apodemes were moderately slender, elongate, darker than the abdominal sternum, rounded and slightly hooked at the apices and subparallel to each other. The abdominal

Figure. 1. Caudal view of second abdominal sternites of Spartina alterniflorasap- feeders. A. Delphacodes penedetecta. B. Megamelus n. sp. C. Prokelisia dolus D. P. marginata. Scales = 0.1 mm. 168 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

h Spartina alterniflora 3.374* ap dept 0.06 ± 0.013 0.05 ± 0.018 0.04 ± 0.013 0.08 ± 0.023 6.961* Sternum 0.03 ± 0.013 0.05 ± 0.016 0.02 ± 0.003 0.06 ± 0.015 mid depth

width 27.378* 0.48 ± 0.051 0.70 ± 0.012 0.47 ± 0.037 0.61 ± 0.059

14.132* apices 0.29 ± 0.025 0.37 ± 0.015 0.27 ± 0.024 0.34 ± 0.031 ., Pd = Prokelisia dolus, Pm marginata. n. sp 50.890* length Apodeme 0.17 ± 0.019 0.06 ± 0.013 0.24 ± 0.014 0.09 ± 0.037

< 0.05. P bases 75.145* 0.23 ± 0.024 0.36 ± 0.011 0.26 ± 0.013 0.35 ± 0.013 < 0.05). P

Table 2. Mean (± standard deviation) tymbal measurements (mm) and analysis of variance for four sap-feeders on (df = 3, 15; Dp Mn Pd Pm F Dp = D. penedetecta, Mn Megamelus * = significant at 2012 THE GREAT LAKES ENTOMOLOGIST 169 sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was short and sinuate on the dorsal aspect and rounded laterally. Megamelus lobatus (Fig. 2C, Table 3). The tymbal apodemes were slender, elongate, darker than the abdominal sternum, rounded at the apices, and diverged toward the apices. The abdominal sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was sinuate on the dorsal aspect and broadly rounded laterally. Neomegamelanus spartini (Fig. 2E, Table 3). The tymbal apodemes were slender, elongate, darker than the abdominal sternum, rounded at the apices, and diverged slightly toward the apices. The abdominal sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was sinuate on the dorsal aspect and rounded laterally. Neomegamelanus elongatus (Fig. 2F, Table 3). The tymbal apodemes were slender, longer than those of N. spartini, darker than the abdominal sternum, rounded at the apices, and subparallel to each other. The abdominal sternum narrowed significantly between the apodemes and the portion extending laterally from the base of each apodeme was convex on the dorsal aspect and rounded laterally. Tumidagena minuta (Fig. 2G, Table 3). The tymbal apodemes were slender, very elongate, darker than the abdominal sternum, rounded at the apices, and curved slightly toward each other. The abdominal sternum narrowed strongly between the apodemes and the portion extending laterally from the base of each apodeme was concave on the dorsal aspect and rounded laterally. Tumidagena terminalis (Fig. 2H, Table 3). The tymbal apodemes were slender, very elongate, darker than the abdominal sternum, rounded at the apices, subparallel and the apices curved sharply toward each other. The abdominal sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was sinuate on the dorsal aspect and rounded laterally. Aphelonema simplex (Fig. 2D, Table 3). The tymbal apodemes of specimens found on S. patens and S. pectinata were very similar and were represented by strongly swollen darkend areas on the dorsal aspect of the abdominal sternum. The abdominal sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was broadly concave on the dorsal aspect and rounded laterally. Amplicephalus simplex (Fig. 2A, Table 3). The tymbal apodemes appeared to be represented by small swollen areas near the middle of the abdominal sternum on the dorsal aspect. The abdominal sternum narrowed strongly between the apodemes and the portion extending laterally from the base of each apodeme was broadly concave on the dorsal aspect and bifurcate laterally. Neohecalus lineatus. See Descriptions of the tymbals of the sap-feeding guild of Spartina pectinata, below. Descriptions of the tymbals of the sap-feeding guild of Spartina pectinata. Anotia fitchi. The male and female specimens examined did not appear to possess tymbals. Megamelus metzaria (Fig. 3A, Table 4). The tymbal apodemes were slender, elongate, darker than the abdominal sternum, subparallel and rounded and slightly hooked at the apices. The abdominal sternum narrowed slightly between the apodemes and the portion extending laterally from the base of each apodeme was convex on the dorsal aspect and rounded laterally. 170 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Figure 2. Caudal view of second abdominal sternites of Spartina patens sap-feeders. A. Amplicephalus simplex B. Delphacodes detecta. C. Megamelus lobatus. D. Aphelo- nema simplex. E. Neomegamelanus spartini. F. N. elongatus. G. Tumidagena minuta. H. T. terminalis. Scales = 0.1 mm.

Prokelisia crocea (Fig. 3B, Table 4). The tymbal apodemes were slender, very elongate, darker than the abdominal sternum, rounded at the apices, and converged toward the apices. The abdominal sternum narrowed strongly between the apodemes and the portion extending laterally from the base of each apodeme was sinuate on the dorsal aspect and rounded laterally. Neohecalus lineatus (Fig. 3C, Table 4). The tymbal apodemes of specimens found on Spartina patens were represented by slightly swollen areas on the dorsal aspect of the abdominal sternum. The abdominal sternum narrowed between the apodemes and the portion extending dorsolaterally from the base of the apodemes was broadly concave on the dorsal aspect and narrowed apically. The tymbal apodemes from specimens found on Spartina pectinata were similar to those from Spartina patens, however, the apodemes were slightly longer. 2012 THE GREAT LAKES ENTOMOLOGIST 171 = Ams , (df = 8, Spartina patens 1.704 Aphelonema simplex ap depth 0.06 ± 0.015 0.05 ± 0.013 0.06 ± 0.000 0.07 ± 0.009 0.07 ± 0.007 0.08 ± 0.000 0.07 ± 0.006 0.07 ± 0.007 0.07 ± 0.007 = Aps

8.030* Sternum T. terminalis, 0.05 ± 0.011 0.04 ± 0.001 0.04 ± 0.000 0.05 ± 0.018 0.02 ± 0.009 0.08 ± 0.000 0.07 ± 0.005 0.07 ± 0.006 0.06 ± 0.017 mid depth = Tt ,

T. minuta width 20.718* = 0.62 ± 0.027 0.52 ± 0.014 0.42 ± 0.025 0.65 ± 0.053 0.46 ± 0.047 1.24 ± 0.030 0.50 ± 0.006 0.75 ± 0.014 0.83 ± 0.016 Tm ,

N. elongatus 26.916* apices = 0.38 ± 0.036 0.19 ± 0.006 0.24 ± 0.030 0.41 ± 0.085 0.14 ± 0.019 0.78 ± 0.013 0.14 ± 0.036 0.15 ± 0.003 0.25 ± 0.016 Ne , .

N. spartini = length 103.765* Apodeme Ns 0.21 ± 0.023 0.14 ± 0.009 0.18 ± 0.008 0.27 ± 0.033 0.15 ± 0.020 0.04 ± 0.000 0.23 ± 0.023 0.04 ± 0.000 0.03 ± 0.007 , N. lineatus = Nl , M. lobatus

= < 0.05. P Ml bases , 597.213* 0.33 ± 0.022 0.18 ± 0.006 0.21 ± 0.017 0.37 ± 0.025 0.14 ± 0.019 0.78 ± 0.013 0.24 ± 0.012 0.15 ± 0.003 0.25 ± 0.016 D. detecta < 0.05).

Table 3. Mean (± standard deviation) tymbal measurements (mm) and analysis of variance for nine sap-feeders on 36; Dd Ml Ns Ne Tm Tt Aps Ams Nl F Dd Amplicephalus simplex * = significant at 172 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Morphometric analyses of tymbals. Analyses of the six features associated with the tymbals for the sap-feeders found on S. alterniflora, S. patens, and S. pectinata resulted in significant differences among the species on each host. All six features, distance between the apodeme bases, apodeme length, distances between the apodeme apices and sternum width, depth at the middle, and depth under the apodeme bases, were significantly different among the four species from S. alterniflora (Table 2). All features, except depth under the apodeme bases, were significantly different among the nine species fromS. patens (Table 3). Four features, except sternum width and depth under the apodeme bases, were significantly different among the four species fromS. pectinata (Table 4). The Hutchinsonian ratio of 1.2:1 to 1.4 :1 has been used as a comparative measure of body size, mass, or length of feeding structures that would allow coexistence among species with similar niches (Hutchinson 1959, Brown 1975, Denno 1980). Comparison of the ratios of the dry body mass of S. patens sapfeeders was used by Denno (1980) to examine niche differentiation among those species. Following Denno (1980), I compared the apodeme lengths of the species on each of the Spartina species. Every one of the species on S. alterniflora differed from the closest species in apodeme length by a ratio > 1.3:1 (Table 5, Fig. 4). Among the S. patens fauna, D. detecta differed from M. lobatus and all others with shorter apodeme lengths, T. terminalis differed from N. elongatus and all others with shorter apodeme lengths, M. lobatus differed from T. minuta and all others with shorter apodeme lengths, N. elongatus differed from N. spartini and the others with shorter apodeme lengths, and T. minuta and N. spartini differed from Am. simplex and Ap. simplex (Table 6, Fig. 5). Among the S. pectinata fauna the two delphacids, P. crocea and M. metzaria had apodeme lengths > 1.3:1 compared to those of Ap. simplex and N. lineatus (Fig. 6).

Discussion Niche differentiation via resource partitioning has been suggested as a result of competition among similar organisms for resources which can include food, nutrients, space, and mates. Numerous studies of vertebrate and invertebrate animals and plants indicate that physical and behavioral attributes of potentially competing species serve to aid in allowing coexistence. This differentiation can

Figure 3. Caudal view of second abdominal sternites of Spartina pectinata sap-feeders. A. Megamelus metzaria. B. Prokelisia crocea. C. Neohecalus lineatus. Scales = 0.1 mm. 2012 THE GREAT LAKES ENTOMOLOGIST 173 (df = Spartina pectinata 1.913 ap depth 0.08 ± 0.013 0.07 ± 0.007 0.08 ± 0.000 0.07 ± 0.003 1.255 Sternum 0.07 ± 0.013 0.06 ± 0.017 0.08 ± 0.001 0.07 ± 0.005 mid depth

width 509.880* 0.83 ± 0.016 1.24 ± 0.030 0.74 ± 0.011 0.58 ± 0.035

94.840* apices 0.28 ± 0.094 0.25 ± 0.016 0.78 ± 0.013 0.46 ± 0.026 65.760* length Apodeme 0.27 ± 0.059 0.03 ± 0.007 0.04 ± 0.000 0.22 ± 0.012

< 0.05. P bases 790.888* 0.29 ± 0.014 0.25 ± 0.016 0.78 ± 0.013 0.40 ± 0.024 < 0.05). P

Table 4. Mean (± standard deviation) tymbal measurements (mm) and analysis of variance for four sap-feeders on 3, 15; Mm Pc Aps Nl F Mm = M. metzaria, Pc P. crocea, Aps Aphelonema simplex, Nl Neohecalus lineatus. * = significant at 174 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Figure 4. Mean tymbal apodeme lengths (mm) of Spartina alterniflora sap feeders. Arcs represent Hutchinsonian ratios > 1.3.

Figure 5. Mean tymbal apodeme lengths (mm) of Spartina patens sap feeders. Arcs represent Hutchinsonian ratios > 1.3. 2012 THE GREAT LAKES ENTOMOLOGIST 175

Figure 6. Mean tymbal apodeme lengths (mm) of Spartina pectinata sap feeders. Arcs represent Hutchinsonian ratios > 1.3. include the partitioning of airborne sounds (Gogala and Riede 1995, Riede and Kroker 1995, Sueur 2002). Niche differentiation among the well-studied sap-feeding guild of leafhoppers and planthoppers on S. patens centered on the stratum of the plant occupied by each species, elevation of the marsh where the hosts of each species occurred, and body size of the insects (Denno 1980). There was little difference in phenology among the species in each stratum suggesting that most of the species are likely to be reproducing during the same time of the season. Thus, it is possible that several species produce substrate-borne vibrational signals on the same plants at the same time. In the leafhopper Graminella nigrifrons (Forbes), the reactions of conspecifics to vibrational signals included alternation of signals which likely resulted from inhibition upon detecting a signal (Hunt and Morton 2001). In the treehopper Ennya chrysura (Fairmaire), adults can produce “jamming” signals that serve to interfere with the courtship of other individuals (Miranda 2006). Artificially produced disruptive vibrational signals have been used to interfere with mating in the leafhopper Scaphoideus titanus Ball (Eriksson et al. 2012). A cacophony of similar signals produced by a variety of species on a plant could potentially reduce reproductive success. Of the guilds of sap-feeding insects examined here, S. patens harbors the greatest number of species, possibly because the habitat in which this grass occurs suffers from less disturbance than that of S. alterniflora (Denno and Roderick 1990). Five of the six analyzed morphometric features were significantly different among the species and the Hutchinsonian ratios for apodeme lengths were > 1.3:1 for each of the species except T. minuta and N. spartini, which occupy different strata of their host plant, and Am. simplex and Ap. simplex, which differ in their ability to colonize new hosts, and N. lineatus, which is rarely collected (Denno 1980). The five species fromS. alterniflora differed in four of the analyzed features and the Hutchinsonian ratios of apodeme lengths were > 1.3:1 for all species combinations. This grass is often subject to more disturbance and is more 176 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 values) for four sap- 1.46 3.13 0.26 2.87 1.20 4.33* q ap depth 1.57 2.12 2.32 4.45* 3.89 6.02* Sternum mid depth

1.52 5.30* 28.64* 23.24* 27.12* 21.82* width

1.52 2.20 6.57* 4.37* 8.09* 5.90* apices ., Pd = Prokelisia dolus, Pm marginata. n. sp 3.25 6.65* 6.24* 9.49* 12.89* 16.14* length Apodeme (df = 12).

< 0.05. 3.14 0.69 P 16.76* 16.07* 13.61* 12.92* bases Spartina alterniflora

Table 5. Pairwise comparisons of mean tymbal measurements (mm) using Tukey’s Honestly Significant Difference ( feeders on Dp - Mn Dp - Pd Dp - Pm Mn - Pd Mn - Pm Pd - Pm Dp = D. penedetecta, Mn Megamelus * = significant at 2012 THE GREAT LAKES ENTOMOLOGIST 177 values) for nine sap- 2.59 2.13 4.32 0.58 1.73 3.00 3.86 3.00 5.18* 2.59 2.42 6.05* 3.46 4.61 5.18* 2.02 2.59 0.86 1.73 2.71 0.29 0.86 4.90 2.30 2.19 0.40 0.58 q ap depth 0.14 0.10 3.19 3.33 7.60* 4.14 2.88 4.31 4.17 2.63 7.44* 7.58* 5.92* 6.17* 6.06* 6.32* 7.50* 4.24 1.24 2.73 2.98 4.53 4.67 3.29 3.44 1.34 10.23* mid depth

6.79* 2.31 9.19* 1.99 8.88* 3.49 3.30 8.36* 1.49 5.30* 6.57* 6.87* 10.36* 15.98* 41.48* 48.35* 55.14* 15.76* 22.55* 12.67* 39.17* 14.37* 10.68* 21.24* 28.03* 13.66* 12.05* width

2.84 2.14 0.02 9.96* 2.46 5.30* 2.44 2.08 5.28* 4.92* 6.91* 3.75 0.91 7.81* 9.05* 12.80* 13.11* 22.58* 12.73* 13.09* 33.23* 30.39* 15.26* 20.43* 15.24* 14.87* 10.65* apices 4.45 6.79* 9.65* 9.62* 6.42* 0.86 3.60 3.22 6.05* 3.57 7.28* 2.83 14.07* 19.26* 19.26* 11.99* 16.44* 10.50* 11.99* 16.44* 13.21* 26.05* 19.98* 26.05* 12.70* 17.15* 26.77* length Sternum (df = 12).

3.08 5.37* 5.41* 8.50* 6.38* 4.56 3.30 7.65* 9.50* 8.33* 5.24* 23.19* 11.80* 23.24* 20.11* 55.78* 22.39* 11.44* 73.61* 70.53* 28.60* 50.42* 16.81* 27.75* 17.82* 14.74* 14.86* bases Apodeme Spartina patens

Table 6. Pairwise comparisons of mean tymbal measurements (mm) using Tukey’s Honestly Significant Difference ( feeders on Dd - Ml Dd - Ns Dd - Ne Dd - Tm Dd - Tt Dd - Aps Dd - Ams Dd - Nl Ml - Ns Ml - Ne Ml - Tm Ml - Tt Ml - Aps Ml - Ams Ml - Nl Ns - Ne Ns - Tm Ns - Tt Ns - Aps Ns - Ams Ns - Nl Ne - Tm Ne - Tt Ne - Aps Ne - Ams Ne - Nl Tm - Tt 178 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 0.86 0.00 1.44 0.86 0.58 0.29 1.15 0.29 0.29 ap depth 1.52 1.26 0.25 8.84* 2.91 1.39 1.64 11.75* 10.48* mid depth 5.48* 51.84* 19.24* 49.85* 32.60* 17.25* 24.73* 27.11* 22.73* width 0.38 0.36 6.20** 6.18* 5.82* 35.69* 35.67* 35.31* 29.48* apices 0.00 0.71 0.71 12.84* 12.84* 22.49* 22.49* 13.56* 23.20* length Sternum

< 0.05. 0.85 1.94 P 79.02* 67.23* 78.17* 10.95* 13.74* 65.28* 12.89* bases Apodeme

Table 6. Continued. Tm - Aps Tm - Ams Tm - Nl Tt - Aps Tt - Ams Tt - Nl Aps - Ams Aps - Nl Ams - Nl Dd = D. detecta, Ml M. lobatus, Ns N. spartini, Ne elongatus, Tm T. minuta, Tt terminalis, Aps Aphelonema simplex, Ams Amplicephalus simplex, Nl = N. lineatus. * = significant at 2012 THE GREAT LAKES ENTOMOLOGIST 179 variable in height than S. patens. The congeners, P. dolus and P. marginata exhibited the greatest amount of difference in tymbal structure compared to other members of the genus (Miller and Wilson 1996). The four species from S. pectinata differed in only two of the analyzed features; Hutchinsonian ratios of apodeme lengths were > 1.3:1 for the two delphacids, P. crocea and M. metzaria versus the caliscelid Ap. simplex and the cicadellid N. lineatus. This tallgrass prairie species is found in much smaller patches than the other Spartina species and is likely fed upon by more species of itinerant generalist leafhoppers (Holder 1990, Wilson et al. 1993). Further, P. crocea is abundant throughout the growing season in the upper portions of this relatively tall grass whereas M. metzaria is found at the base of the plants and is only commonly collected in the early fall (Holder and Wilson 1992). The caliscelid Ap. simplex and leafhopper N. lineatus are usually very rare. At one location, Blue Lick Natural Area, Pettis County, Missouri, both N. lineatus and M. metzaria were enormously abundant and P. crocea was absent (Wilson, personal observation). Host plants provide resources for sap-feeding insects that include nutri- tion, water, oviposition sites, and a substrate for communication via vibrational signals. The partitioning of resources has been shown to allow coexistence of potentially competing species (Schoener 1974, McClure and Price 1975, Denno 1980). The partitioning of food resources has been related to differences in the morphology of structures associated with feeding, such as beak size in finches (Grant and Grant 2006) and proboscis length in bumblebees (Pyke 1982). Tymbal morphology likely affects the vibrational signals used to recognize conspecifics. If so, then the occurrence of several species on the same host could result in a cacophony of signals unless the “substrate resource” is partitioned in some man- ner. Tymbal morphology could be indicative of reproductive isolation among species and may be useful for determining sibling species as well as providing insights about behavior, ecology, and evolution in guilds of sap-feeding insects that communicate via substrate vibrations.

Acknowledgments It is with great pleasure that I dedicate this paper to Dr. J. E. “Jay” McPherson, my Ph. D. advisor, who taught, above all, that insects are fascinat- ing creatures and an understanding of the natural history of a group requires an entomologist to be a taxonomist, morphologist, and ecologist. Thanks are due to Ms. Amy Miller, Department of Biology, University of Central Missouri, Warrensburg, for helping take the measurements and draw pencil illustrations. I would also extend my deep appreciation to the late Dr. Robert F. Denno, De- partment of Entomology, University of Maryland, College Park, for donating several specimens used in this study and for his helpful comments on an earlier draft of this paper.

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Whitcomb, R. F., J. Kramer, J. E. Coan, and A. L. Hicks. 1987. Ecology and evolution of leafhopper-grass host relationships in North American grasslands. In K. F. Harris (ed.), Current topics in vector research 4: 121–178. Wilson, S. W., J. L. Smith, and P. D. Calvert. 1993. Planthoppers of a Missouri tall-grass prairie (Homoptera: Fulgoroidea). Journal of the Kansas Entomological Society 66: 75–80. Zezlina, I., and M. Virant-Doberlet. 2007. Vibrational communication of Metcalfa pruinosa (Hemiptera: Fulgoroidea: Flatidae). Annals of the Entomological Society of Amererica 100: 73–82. Zhang, Z. T., W. Z. Korg, J. L. Gao, and T. H. Shao. 1988. Acoustic signal-producing organ of brown planthopper (BPH). International Rice Research Newsletter 13: 38–39. 2012 THE GREAT LAKES ENTOMOLOGIST 183 Pentatomoidea (Hemiptera: Pentatomidae, Scutelleridae) Associated with the Dioecious Shrub Florida Rosemary, Ceratiola ericoides (Ericaceae) A. G. Wheeler, Jr.1

Abstract Florida rosemary (Ceratiola ericoides), an ericaceous shrub with needlelike leaves, is characteristic of Florida scrubs and certain other xeric plant communities of well-drained sands. The plant is dioecious, its inconspicuous male and female flowers borne on separate individuals. From 2003 to 2012 (mainly 2007–2012), rosemary was sampled periodically (with all months included at least once during the period) in sand ridges of peninsular Florida and two riverine dunes in southeastern Georgia. Branches of male and female plants were tapped separately into a beating net. Sampling during the final three years was female-plant-biased to facilitate work on fruit- and seed-feeding heteropterans. Nymphs and adults of three pentatomoid species were found on C. ericoides: the pentatomid Thyanta custator custator (F.) and scutellerids Diolcus chrysorrhoeus (F.) and Homaemus proteus Stål. Only T. c. custator was taken in both states, occurring at 19 sites (19 nymphs, 53 adults); D. chrysorrhoeus was found in Florida at 16 sites (9 nymphs, 165 adults). The collection of T. c. custator and D. chrysorrhoeus from female rosemary plants essentially throughout the sampling period, including nymphs, exuviae, and mating pairs, coupled with their near absence from male plants, suggests that the bugs are not incidental on rosemary but feed on its fruits. Whether either species com- pletes its life cycle on rosemary is unknown. Three nymphs and eight adults of the little-known H. proteus were collected from female rosemary plants at four sites, but the scutellerid’s relationship to C. ericoides remains to be determined. Briefly noted is the collection of the pentatomid Euschistus obscurus (Palisot de Beauvois) and scutellerid Stethaulax marmoratus (Say), whose adults were collected infrequently on female rosemary plants in Florida.

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Florida rosemary (or sand heath; hereafter rosemary), Ceratiola ericoides L. (Ericaceae; formerly Empetraceae), is a characteristic plant of Florida scrubs (Mulvania 1931, Austin et al. 1987, Menges and Kohfeldt 1995). These plant communities are xeric, pyrogenic, shrub-dominated, and occur on excessively well-drained deep sands of ancient shorelines and ridges (Laessle 1958, Myers 1990, Menges 1999). Although biodiversity of scrubs is lower than for certain other plant communities, these ecological islands harbor numerous endemic (precinctive) and near-endemic animals and plants, many of them threatened or endangered (Deyrup 1989, Christman and Judd 1990, Menges 1999, Lamb et al. 2006, FNAI 2010). Scrubs were widespread in Florida during the late Pliocene and early to mid-Pleistocene before fragmentation into patches with a shift from xeric to mesic conditions (Webb 1990, Branch et al. 2003). Especially since 1900, suburban and urban development and the clearing of land for citrus and other crops have reduced the extent of scrub communities (Huck et al. 1989,

1School of Agricultural, Forest, and Environmental Sciences, Clemson University, Clemson, SC 29634-0310. (e-mail: [email protected]). 184 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Christman and Judd 1990, Myers 1990). In addition, fire suppression can be detrimental to scrub biodiversity (Menges 1999). Rosemary in Florida is locally abundant in scrubs of the Mt. Dora Ridge (Ocala National Forest), Lake Wales Ridge, and certain other sand ridges of the central peninsula. It also is common in scrubs of the Atlantic Coastal ridge and Panhandle coast, and persists elsewhere in smaller patches. Rosemary scrubs (balds), dominated by C. ericoides, are a community variant found on the driest ridge crests, such as in the southern Lake Wales Ridge (e.g., Archbold Biologi- cal Station), and on barrier islands of the Panhandle (FNAI 2010). Ceratiola ericoides ranges north, with disjunct populations in Georgia and South Carolina, and southwest to the Florida Panhandle, southwestern Alabama, and southeastern Mississippi (Abrahamson et al. 1984, Wally and Menges 2002, FNAI 2010, USDA NRCS 2012). Ceratiola ericoides belongs to a monotypic genus traditionally placed in the small family Empetraceae but is now considered a member of a redefined, more inclusive Ericaceae (Kron et al. 2002). The shrub has evergreen needlelike leaves, is wind pollinated, and bears inconspicuous male and female flowers on separate individuals (dioecy). Plants bloom mainly from September to November (Trapnell et al. 2007, 2008; Wheeler 2009), but under certain conditions can flower at other times (Patrick et al. 1995). Fruits are two-seeded drupes (Small 1933, Godfrey 1988), with clusters containing greenish-yellow, fleshy (ca. 2–3 mm) fruits, as well as dry, hard fruits (Schmidt 2008, Trapnell et al. 2008). Fleshy fruits, generally less numerous than dry fruits, are assumed to result from pollinated flowers (Schmidt 2008). Rosemary is aromatic (Hubbell and Walker 1928, Tanrisever et al. 1987, Jordan et al. 1992), presumably well defended against herbivory by insects (Deyrup 1989; Deyrup and Eisner 1993, 2001), and is fed on by relatively few insects (Sandoval-Mojica and Capinera 2011). Ceratiola specialists include a grasshopper, Schistocerca ceratiola Hubbell and Walker (Franz and Franz 1989, Smith and Capinera 2005); geometrid moth, Nemoria outina Ferguson (Deyrup and Eisner 1993, Eisner et al. 2005, Canfield et al. 2008); leafhopper, Alconeura bisagittata (Beamer) (Wheeler and Hicks 2012); and plant bugs Kelto- nia balli (Knight) (Wheeler 2009) and Parthenicus weemsi Henry (Henry 1982). The relatively few generalist herbivores of rosemary include the walkingstick Anisomorpha buprestoides (Stoll) (Conle et al. 2009), scale insects such as the coccid Neopulvinaria innumerabilis (Rathvon) (Hamon and Williams 1984) and diaspidid Rhizaspidiotus dearnessi (Cockerell) (Beshear et al. 1973), and two rhyparochromid seed bugs (A.G.W., unpubl. data). Here I document the association of three pentatomoid heteropterans with C. ericoides: the scutellerids Diolcus chrysorrhoeus (F.) and Homaemus proteus Stål, and the pentatomid Thyanta custator custator (F.). I take pleasure in dedi- cating this paper to J. E. McPherson, an esteemed friend and colleague who has contributed substantially to our knowledge of the bionomics and systematics of North American Pentatomoidea.

Methods and Study Sites Pentatomoid heteropterans were collected periodically, beginning in 2003 but mainly from 2007 to 2012, during studies of other rosemary insects (Wheeler 2009, Wheeler and Hicks 2012). Ceratiola ericoides was sampled at irregular intervals, primarily in peninsular Florida scrubs such as the Crescent City, Lake Wales, and Northern Brooksville ridges, and in two riverine dunes of the Ohoopee Dunes system in the coastal plain of southeastern Georgia. Male and female plants were sampled separately by using an ax handle to tap branches over a short-handled beating net with a shallow bag. From 2010 to 2012, sampling was biased toward female plants to increase the probability 2012 THE GREAT LAKES ENTOMOLOGIST 185 of encountering fruit-feeding pentatomoids. Sampling time depended on the size of a rosemary population and ranged from about 10 min (< 5 plants) to 45 min (> 50 plants). A woody vine, earleaf greenbrier (Smilax auriculata Walt.; Smilacaceae), which sometimes grew over rosemary plants, was avoided during sampling except when greenbrier fruits were sampled for possible individuals of Pentatomoidea. Nymphs and adults of pentatomids and scutellerids from rosemary were collected into snap-cap plastic vials, and nymphs later were sorted to instar using a stereoscopic microscope. Several fifth instars ofD . chrysorrhoeus and T. c. custator were held with sprigs of rosemary to obtain adults for species identifications. Late instars assumed to be those ofH. proteus were not reared to adulthood; they tentatively were identified based on their collection from female plants that yielded adults of H. proteus, smaller size compared to late instars of D. chrysorrhoeus, and head structure more similar to that of Homaemus than Diolcus. Nymphs thought to represent H. proteus also resembled those of H. variegatus Van Duzee taken on the rosaceous shrub chamise, Adenostoma fasciculatum Hook. and Arn., in California (A.G.W., pers. obs.). In listing specimens examined, Roman numerals are used to indicate nymphal instars and are preceded by Arabic numerals referring to the numbers collected for each instar. The number of males and females is given, or number of adults when gender is unknown. Voucher material has been deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM).

Results Diolcus chrysorrhoeus (F.) This pachycorine scutellerid was described from “Carolina” by Fabricius (1803); Scutelleria viridipunctata, described from Florida by Say (1832), was synonymized with D. chrysorrhoeus by Uhler (1859). Common in Florida, where it has been recorded from Jacksonville and St. Augustine in the north (Barber 1914) to the Everglades and Keys in the south (Blatchley 1926, Torre-Bueno 1933), D. chrysorrhoeus ranges west to Arkansas and eastern Texas. An Illinois record is of questionable validity (McPherson 1982); a provisional record from Virginia was published by Hoffman (1994). Ecological data on this scutellerid are scant. In swamps near Mansura in central Louisiana, ten adults were collected from Spanish moss, Tillandsia usneoides (L.) L. (Bromeliaceae), in June 1909, but none were found during more extensive sampling of the moss during the winter of 1908–1909 (Rosenfeld 1911). Blatchley (1926), however, noted that adults overwinter in clumps of Spanish moss in Florida, and Rainwater (1941) collected D. chrysorrhoeus from Span- ish moss near Tallulah in northeastern Louisiana. Adults in Florida are found on trees and shrubs (Blatchley 1926) and have been recorded from holly, Ilex sp. (Aquifoliaceae), and observed feeding on developing fruits of guava (Lattin 1964), presumably the introduced Psidium guajava L. This scutellerid has been collected in South Carolina (Sullivan’s Island) from common wax-myrtle or southern bayberry, Morella cerifera (L.) Small (Myricaceae) (Krauss 1964). I have taken nymphs and adults of D. chrysorrhoeus on fruits of M. cerifera at Charleston, South Carolina, in late October 2008 (USNM collection). Overgaard (1968) reported D. chrysorrhoeus as a possible predator of the southern pine beetle, Dendroctonus frontalis Zimmermann, in Texas, but scutellerids are regarded as strict plant feeders (Schuh and Slater 1995, Javahery et al. 2000). Association with Ceratiola ericoides. Diolcus chrysorrhoeus (Figs. 1, 2) was found on fruit-bearing rosemary plants at 16 sites in peninsular Florida, from the Crescent City Ridge of Putnam County south to the southern Lake Wales Ridge in Highlands County. One first instar, one third instar, two fourth instars, five fifth instars, and 165 adults were observed. The largest 186 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Figure 1. Diolcus chrysorrhoeus, third-instar nymph (collected as instar I), Levy County, Florida, Rt. 27A, 3.7 km SE of Bronson, 18 August 2012.

Figure 2. Diolcus chrysorrhoeus, adult, collected with the first-instar nymph in Levy County, Florida. 2012 THE GREAT LAKES ENTOMOLOGIST 187 numbers of adults in any sample were 56 in Levy County in early June and 30 in Gilchrist County in late March. Nymphs were found on rosemary in early June, mid-August, and from late October to late December; adults, from late January to mid-August and late October to late December. A mating pair was observed in mid-June. Specimens examined. FLORIDA: Alachua Co., Rt. 24, 1.8 km SW of Archer, 29°31.585΄N 82°32.079΄W, 9 June 2012, 1♂; Rt. 24, 3.5 km SW of Archer, 29°30.928΄N 82°33.046΄W, 31 Oct. 2009, 2♂, 1♀ & 29 July 2010, 1♂ & 12 Nov. 2010, 1♂. Gilchrist Co., Rt. 129, 2.6 km N of Bell, 29°46.680΄N 82°51.996΄W, 30 Mar. 2012, 15♂, 15♀ & 9 June 2012, 1–III, 1–V, 2♂, 1♀; Rt. 26, 10.5 km E of Trenton, 29°37.816΄N 82°42.643΄W, 9 June 2012, 1♀. Highlands Co., S of Rt. 621 W of Daffodil St., 7.2 km NW of Lake Placid, 27°19.229΄N, 81˚25.525΄W, 15 Nov. 2003, 1 adult & 27 Nov. 2004, 3 adults; Rt. 70, 8.5 km S of Lake Placid, 27˚12.555΄N, 81˚21.426΄W, 7 Nov. 2007, 1♂, 1♀. Lake Co., Rt. 19, 1.5 km S of Rt. 40 & W of Astor Park, Ocala National Forest, 29°08.008΄N 81°37.039΄W, 7 Nov. 2007, 1-V, 2 adults. Levy Co., Rt. 24, 1.5–1.6 km S of Alachua Co. line SW of Archer, 29°30.252–254΄N 82°34.181–186΄W, 31 Oct. 2009, 1–V, 2♂, 3♀ & 14 Mar. 2012, 1♂; Rt. 27A, 3.7 km SE of Bronson, 29°25.920΄N 82°36.363΄W, 31 Oct. 2009, 1–IV & 31 Mar. 2012, 4♂, 4♀ & 9 June 2012, 1–IV, 37♂, 19♀(including mating pair) & 18 Aug. 2012, 1–I, 18♂, 8♀; Rt. 27A, 4.2 km SE of Bronson, 29°25.906΄N 82°36.327΄W, 14 Mar. 2012, 3♂, 1♀; NE 62nd St., 0.4 km N of Rt. 27A, 6.5 km SE of Bronson, 29°25.603΄N 82°34.892΄W, 31 Mar. 2012, 2♂, 1♀; Rt. 27A, 10 km SSE of Bronson, 29°24.729΄N 82°32.686΄W, 9 Nov. 2008, 4♂ & 22 Dec. 2011, 1♂ (dead when collected) & 14 Mar. 2012, 1♀ & 18 Aug. 2012, 1♂. Marion Co., Rt. 40, Ocala National Forest, 16 km E of Lynne, 29°10.632΄N 81°46.573΄W, 23 Dec. 2011, 1–V (molted to adult ♂), 1♂, 2♀; For. Rd. 13, Ocala National Forest, 0.6 km E of Rt. 40, 11 km W of Juniper Springs, 29°10.352΄N 81°45.379΄W, 21 Jan. 2012, 2♂. Putnam Co., jct. Rt. 20 & Royal Way, 5.7 km W of Interlachen, 29°37.533΄N 81°56.752΄W, 18 Aug. 2012, 1♂, 1♀; Johnson, nr jct. Co. Rd. 20A & Rt. 21, 29°35.526΄N 81°58.532΄W, 22 Nov. 2009, 1–V. Homaemus proteus Stål The pachycorine scutellerid H. proteus was described by Stål (1862) from Mexico and now is known to occur south to Colombia. U.S. records mostly are western: California, New Mexico, and Texas (Froeschner 1988), as well as Ari- zona (Van Duzee 1917) and Nevada (Christensen 1970). Homaemus proteus was listed from Iowa by Osborn (1890), but Van Duzee (1904) questioned the record, as did Stoner (1920), who noted that all other U.S. records are southwestern. Neither Barber (1914) nor Blatchley (1926) recorded this scutellerid from Florida. Walley (1929) was the first to report it from the state: “eastern Florida (Ashmead)”; the record apparently is based on a specimen (♀) in the USNM collection, labeled “So. Fla./Ashmead USNM Acc. 18050.” W. H. Ashmead, a hy- menopterist, donated his collection to the USNM in 1898 (USNM 1976); insects of orders other than Hymenoptera, including the specimen of H. proteus, probably were part of the donation. Froeschner (1988) included Florida in the U.S. distribution, but H. proteus was not included in Eger and Baranowski’s (2002) key to Florida Scutelleridae despite recent collections from Florida. The first Florida record published since Walley’s (1929) paper was from Alachua County (Dixon and Anderson 2005). Additional Florida county records are Charlotte, Lee, Orange, Palm Beach, and Pinellas (Dixon and Anderson 2006–2010), as well as Hernando, Hillsborough, Martin, and Miami-Dade (I. C. Stocks, pers. comm.). Host relationships for H. proteus are uncertain because collection records do not mention the presence of nymphs. Torre-Bueno (1939) mentioned its occurrence on, or near, the roots of grasses. Such habits might characterize the more frequently encountered and widespread H. parvulus (Germar) (Slater and Baranowski 1978, McPherson 1982, Jones 1993, A.G.W., pers. obs.), but probably do not apply to H. proteus. Recent records of H. proteus mainly are from 188 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 composites (Asteraceae): Parthenium argentatum Gray in Texas (Stone and Fries 1986), P. hysterophorus L. in Mexico (McClay et al. 1995), and Ambrosia artemisiifolia L. (Dixon and Anderson 2007, 2010) and Helianthus sp. (Dixon and Anderson 2005) in Florida. Additional collection records include Lantana camara L. (Verbenaceae) in Mexico (Palmer and Pullen 1995) and Gaura angustifolia Michx. (Onagraceae) in Florida (Dixon and Anderson 2006). Association with Ceratiola ericoides. Homaemus proteus was collected from female rosemary plants at four sites in peninsular Florida. Five of the eight adults and the only nymphs (1 fourth and 2 fifth instars) were found at a site in Putnam County (Crescent City Ridge). Single adults were collected in scrubs of Alachua and Levy counties (Northern Brooksville Ridge) and Highlands County (southern Lake Wales Ridge). Specimens examined. FLORIDA: Alachua Co., Rt. 24, 3.5 km SW of Archer, 29°30.928΄N 82°33.046΄W, 29 July 2009, 1 adult. Highlands Co., 8.5 km S of Lake Placid, 27°12.555΄N, 81°21.426΄W, 23 Mar. 2007, 1♂. Levy Co., Rt. 27A, 3.7 km SE of Bronson, 29°25.920΄N 82°36.363΄W, 9 June 2012, 1♂. Putnam Co., jct. Rt. 20 & Royal Way, 5.7 km W of Interlachen, 29°37.533΄N 81°56.752΄W, 23 Jan. 2010, 1♀ & 15 Apr. 2010, 1♀ & 19 June 2010, 1–V, 1♂ & 20 Jan. 2012, 1♀ & 18 Mar. 2012, 1–IV & 8 June 2012, 1–V, 1♂. Thyanta custator custator (F.) Fabricius (1803) described Cimex custator from “Carolina”; Stål (1862) included the species in his new genus Thyanta. The original description was sufficient to fix the species, but subsequent workers often identified other large (at least 8 mm) species of the genus as T. custator, even though specimens did not conform to the Fabrician description and were from other regions of North America. The confused taxonomic history of T. custator and similar taxa largely was resolved by Ruckes (1957), who regarded the nominate form as stable morphologically and restricted to the Atlantic and Gulf coasts (inland to ca. 120 km) from Massachusetts (since deleted from the range; Rider and Chapin 1992) and New York (Long Island) south to Florida and west to central Louisiana. The record from extreme southern Ohio (Ruckes 1957: p. 7, footnote 1) is based on a misidentification ofT . calceata (Say) (D. A. Rider, pers. comm.). Ruckes (1957) placed the morphologically similar form accerra McAtee as a subspecies of T. pallidovirens (Stål). Rider and Chapin (1992), however, gave accerra subspecific status within T. custator; the range of T. c. accerra, found in the eastern two- thirds of the United States, overlaps only narrowly with that of the nominate subspecies (Rider and Chapin 1992). Most plant associations and biological data attributed to T. c. custator are based on studies outside the range of the nominate subspecies and refer to T. c. accerra (Ruckes 1957, Rider and Chapin 1992, map 2, p. 52). Thus, a biological study of “T. custator” in Missouri (Oetting and Yonke 1971) pertains to T. c. accerra (McPherson 1982), as do host associations in Arizona (Jones 1993) and those of Furth (1974), which were based mainly on records by Hart (1919) and Esselbaugh (1948) for Illinois (McPherson 1979, 1982). Furth (1974) also included under “T. custator” the hosts given by Torre-Bueno (1939), but Torre-Bueno’s concept of T. custator included both the nominate subspecies and accerra (Rider and Chapin 1992). Few host records for T. custator, therefore, can be attributed reliably to the nominate subspecies. An exception, as noted by McPherson (1982), is Dozier’s (1920) Florida record from “haw bush” [Crataegus sp.; Rosaceae]; T. c. accerra is not known from Florida (Rider and Chapin 1992). Because even valid plant records for T. c. custator probably are based only on the collection of adults, host relationships of this subspecies should be considered undocumented. Association with Ceratiola ericoides. Thyanta c. custator was collected from rosemary in Georgia (2 sites: 6 nymphs, 7 adults) and peninsular Florida (17 sites: 13 nymphs, 46 adults). At an Emanuel County site in Georgia, two 2012 THE GREAT LAKES ENTOMOLOGIST 189 mating pairs were observed in mid-March, a third instar and two fifth instars in late April, a third instar in mid-May, and second instars in mid-May and mid-September. Single adults were found in late April, mid-July, and late No- vember. I encountered nymphs in Florida in late March, early June, late October, mid-November, and late December. Adults were collected in late January, late March, early and mid-June, late October, late November, and late December. Mating pairs were found in mid- and late March. Specimens examined. FLORIDA: Alachua Co., Rt. 24, 3.5 km SW of Archer, 29°30.928΄N 82°33.046΄W, 31 Oct. 2009, 2–V, 2♂, 3♀. Clay Co., Rt. 21, 3.2 km NE of Keystone Heights, 29°48.536΄N 82°00.575΄W, 22 Nov. 2009, 6 adults & 23 Jan. 2010, 1♂ & 20 Mar. 2010, 2♂ & 21 Jan. 2011, 1♂, 1♀. Gilchrist Co., Rt. 129, 2.6 km N of Bell, 29°46.680΄N 82°51.996΄W, 31 Oct. 2009, 1 adult & 9 June 2012, 1♂; Rt. 26, 10.5 km E of Trenton, 29°37.816΄N 82°42.643΄W, 9 June 2012, 1–V (molted to adult ♂). Highlands Co., nr jct Lancaster & Livingston Ave, 7 km NW of Avon Park, 27°38.375΄N 81°33.453΄W, 22 Nov. 2009, 1 adult & 13 Nov. 2010, 1–V (reared to adult ♂). Levy Co., Rt. 24, 1.5–1.6 km S of Alachua Co. line SW of Archer, 29°30. 252– 254΄N 82°34.181–186΄W, 31 Oct. 2009, 1 adult & 14 Mar. 2012, 1♂, 1♀; Rt. 27A, 3.7 km SE of Bronson, 29°25.920΄N 82°36.363΄W, 31 Oct. 2009, 1 adult & 23 Jan. 2010, 1 ♂ & 31 Mar. 2012, 2–II; Rt. 27A, 4.2 km SE of Bronson, 29°25.906΄N 82°36.327΄W, 14 Mar. 2012, 2♂, 1♀ (including mating pair); NE 62nd St., 0.1 km N of Rt. 27A, 6.5 km SE of Bronson, 29°25.484΄N 82°34.869΄W, 31 Mar. 2012, 1♂, 2♀ (including mating pair); NE 62nd St., 0.4 km N of Rt. 27A, 6.5 km SE of Bronson, 29°25.603΄N 82°34.892΄W, 31 Mar. 2012, 1–III; Rt. 27A, 10 km SSE of Bronson, 29°24.729΄N 82°32.686΄W, 14 Mar. 2012, 1♂; Co. Rd. 32, 1.8 km N of Bronson, 29°27.825΄N 82°37.770΄W, 31 Oct. 2009, 3♀. Marion Co., Rt. 40, Ocala National Forest, 16 km E of Lynne, 29°10.632΄N 81°46.573΄W, 23 Dec. 2011, 3–V (all molted to adult ♀); Ocala National Forest, 0.6 km E of Rt. 40, 3.4 km W of Juniper Springs, 29°10.663΄N 81°44.568΄W, 21 Jan. 2012, 1♂, 1♀. Putnam Co., jct. Rt. 20 & Royal Way, 5.7 km W of Interlachen, 29°37.533΄N 81°56.752΄W, 23 Jan. 2010, 2♀ & 19 June 2010, 1–III, 2–V, 1 adult & 20 Jan. 2012, 1♀ & 18 Mar. 2012, 1♂, 1♀ (mating pair); Illinois St., 0.1 km E of Co. Rd. 315, 5 km N of Interlachen, 29°40.106΄N 81°53.698΄W, 18 Mar. 2012, 1♂, 2♀; Co. Rd. 315, 2.2 km S of Interlachen, 29°36.531΄N 81°53.186΄W, 23 Dec. 2011, 1♂. GEORGIA: Emanuel Co., Old Hwy. 46, 0.1 km E of Rt. 1, 4.8 km N of Oak Park, 32°23.704΄N 82°18.474΄W, 19 Mar. 2011, 2♂, 2♀ (mating pairs) & 30 Apr. 2011, 1–III, 2–V, 1♀ & 15 May 2011, 1–III & 16 July 2011, 1 adult & 17 Sept. 2011, 1–II & 23 Nov. 2011, 1♀ & 19 May 2012, 1–II; Tattnall Co., Handy Kennedy Rd. (=Co. Rd. 90), 1.3 km N of Rt. 152, 7.7 km W of Cobbtown, 32°17.713΄N 82°11.928΄W, 23 Nov. 2011, 1–V. Other Pentatomoidea Adults of two other pentatomoid species, whose relationship to Ceratiola ericoides might be incidental, also were collected on female rosemary plants. Pentatomidae. In Florida, I collected a female of Euschistus obscurus (Palisot de Beauvois) in Alachua County (SW of Archer) on 31 October 2009, a male in Levy County (3.7 km SE of Bronson) on 18 August 2012, and a female in Putnam County (5.7 km W of Interlachen) on 18 August 2012. Cotton and soybean are among the hosts of E. obscurus in southeastern states (Bundy and McPherson 2000). Scutelleridae. Four males of Stethaulax marmoratus (Say) were collected in Levy County, Florida (3.7 km SE of Bronson), on 9 June 2012, and a female was taken on the same date at a nearby site in Levy County (6.5 km SE of Bronson). Adults are known from plants of several families, including species of Juniperus and Thuja (Cupressaceae) and Rhus (Anacardiaceae). Nymphs have been collected from R. copallina L. and R. glabra L. (Walt and McPherson 1972, 1973; McPherson 1982; Tecic and McPherson 2004). 190 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Discussion Information on host plants of the three pentatomoid species found on Ceratiola ericoides has remained scant, even for the nominate subspecies of the common pentatomid T. custator. In harboring nymphs (mostly late instars) of the scutellerids D. chrysorrhoeus and H. proteus and pentatomid T. c. custator, the shrub, thus, meets an important criterion of host status. Only one or two individuals of H. proteus were collected in any sample, but adults of D. chrysorrhoeus sometimes were numerous on C. ericoides. Moreover, D. chrysorrhoeus and T. c. custator were collected over multiple years and seasons, and mating pairs were observed. All three pentatomoid species were taken almost exclusively on female rosemary plants and appeared not to be associated with Smilax auriculata, a vine that grew on rosemary at certain sites. Because many species of Pentatomidae (Panizzi et al. 2000) and Scutelleridae (Javahery et al. 2000) feed on fruits and seeds, the three species found on rosemary might feed similarly. I did not, however, observe feeding in the field. Assuming the bugs are frugivores on rosemary, is their feeding restricted to fleshy fruits, which often are unavailable or scarce when the pentatomoids are present on the plants, or can they feed on the smaller, dry fruits, as do the rhyparochromid bugs found on rosemary (A.G.W., unpubl. data)? Feeding on the shrub’s vegetative structures also is possible. Evidence suggests that the occurrence of D. chrysorrhoeus and T. c. custator on rosemary is not incidental, but additional study is needed to determine whether they can complete their life cycles on rosemary (and, if so, the number of generations) and to clarify the role that this plant might play in the bugs’ performance and fitness.

Acknowledgments I gratefully acknowledge Thomas J. Henry (Systematic Entomology Labo- ratory, USDA, ARS, Washington, DC) for identifying Pentatomoidea, David A. Rider (Department of Entomology, North Dakota State University, Fargo) for clarifying the status of the Ohio record of Thyanta c. custator and allowing me to refer to the information as a personal communication, Ian C. Stocks (Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville) for providing information on records of Homaemus proteus in Florida, including label data from specimens in the Florida State Collection of Arthropods, and Gerald R. Carner (School of Agricultural, Forest, and Environ- mental Sciences, Clemson University) for photographing Diolcus chrysorrhoeus.

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Abstract A list of the Pentatomidae of New Mexico with county records and collection dates is given. A total of 87 species of stink bugs is reported for New Mexico with 19 new state records.

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Little is known about the stink bugs (Heteroptera: Pentatomidae) of New Mexico. A few partial lists of the Heteroptera, including stink bugs, have been compiled for specific regions in the state (e.g., Uhler 1871, 1876; Townsend 1894; Van Duzee 1903; Uhler 1904; Barber 1926; Stroud 1950). Also, the Catalog of the Heteroptera (Froeschner 1988) is a valuable reference for state records, including New Mexico, although several pentatomid groups have been revised since its publication. However, little information is available on the distribution of stink bugs within New Mexico. The most comprehensive work is that of Ruckes (1937, 1938) who generated a partial list of the pentatomids for the state based on brief field collections from several regions, with an emphasis on northern New Mexico. Presented here is the first attempt to compile a list of the stink bugs of New Mexico with in-state distributions and collection dates. I am pleased to dedicate this paper to Dr. J. E. McPherson in honor of his contributions to the field of pentatomology. He has been a mentor and friend, introducing me to the Heteroptera. His research has greatly increased our understanding of the biology and morphology of the Pentatomidae and other heteropteran fauna of North America.

Materials and Methods Distribution data for each species are based on county records compiled from field-collections, museum specimens, and literature records. Field collections were made from 2000 to 2012 in a variety of habitats, predominantly in southern New Mexico, using several collection methods, including sweep net, beat net, black light, and hand collecting. Collection data from identified adult museum specimens in the Arthropod Collection of New Mexico State Univer- sity in Las Cruces, the Insect Collection of Eastern New Mexico University in Portales, and the Museum of Southwestern Biology Arthropod Collection in Albuquerque were taken. I made the majority of stink bug identifications. Don Thomas identified species for a few problematic groups and also made the majority of identifications from the Portales collection. Distributional data represented here for each species are based on county records, supplemented with data from Ruckes original list (1937, 1938), Froeschner (1988), and others. Location of counties in New Mexico are presented in Figure 1. Collection dates are based on earliest and latest collections of adult bugs. I placed species alphabetically within subfamily to facilitate use of the list. See the online Pentatomoidea Home page for tribal relationships within the Pentatominae (Rider 2009).

1Department of Entomology, Plant Pathology, and Weed Science, New Mexico State University, Las Cruces, NM 88003. 2012 THE GREAT LAKES ENTOMOLOGIST 197

Figure 1. Counties of New Mexico.

Results A total of 87 species of pentatomids are reported for New Mexico with 19 new state records. Checklist of Pentatomidae of New Mexico Subfamily Asopinae Alcaeorrhynchus grandis (Dallas) Collection dates: 15 June 2007 (1 ♀) and “Aug.-Dec.” 2007 (1 ♀). Counties: Otero and Roosevelt. Note: This is the first record of this species from New Mexico. Apateticus lineolatus (Herrich-Schaeffer) Collection dates: 18 September 1975. Counties: Roosevelt. Note: This is the first record of this species from New Mexico. 198 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Apateticus marginiventris (Stål) Collection dates: 20 July 2004 (1 ♂, current study), 21 July 1916 (Barber 1926), and 25 August 1932 (Ruckes 1937). Counties: Colfax, Otero, and Santa Fe. Apoecilus bracteatus (Fitch) Collection dates: 15 July (no year given, 1 individual) (Ruckes 1937), 3-5 August (no year given, as Apateticus crocatus [Uhler]) (Ruckes 1938), and 3 and 11 August (no year given, 3 ♂, 1 ♀) (Uhler 1904). Counties: Bernalillo, San Miguel, and Santa Fe. Note: Froeschner (1988) suggested that this state record needs to be verified. Apoecilus cynicus (Say) Collection dates: 11 and 14 August (no year given) (1 ♂, 1 ♀, respectively) (Uhler 1904). Counties: Santa Fe. Perillus bioculatus (Fabricius) Collection dates: 14 March through 27 October. Counties: Bernalillo, Chaves, Colfax, Doña Ana, Eddy, Grant, Hard- ing, Hidalgo, Lea, Roosevelt, Santa Fe, Taos, Union, and Valencia. Note: County records from Ruckes (1937) are included here. Perillus confluens (Herrich-Schaeffer) Note: Uhler (1876) and Ruckes (1937) listed this species from New Mexico, the latter author could not verify its presence in the state. Perillus exaptus (Say) Collection dates: 25 July (no year given, multiple individuals, Ruckes 1937) and 12 August 1964 (1 ♂, current study). Counties: Colfax and Grant. Perillus splendidus (Uhler) Collection dates: 11 April through 14 September. Counties: Catron, Doña Ana, and Taos. Note: This is the first record of this insect from New Mexico. Perillus strigipes (Herrich-Schaeffer) Collection dates: 28 August, 1932 (two specimens, Ruckes 1937). Counties: Colfax. Podisus maculiventris (Say) Collection dates: 21 March through 23 October. Counties: Bernalillo, Doña Ana, Otero, Roosevelt, Santa Fe, Taos, and Valencia. Note: The only previous record of this species for the state (as P. modestus [Dallas], which has been synonymized with P. maculiventris) is that of Ruckes (1937), county record included here. Podisus placidus Uhler Collection dates: “April” through 4 September. Counties: Doña Ana and Roosevelt. Note: This is the first record of this species from New Mexico. 2012 THE GREAT LAKES ENTOMOLOGIST 199 Podisus serieventris Uhler Collection dates: 28 March through 23 September. Counties: Doña Ana, Lincoln, Otero, and Santa Fe. Note: A county record from Ruckes (1937) is included here. Stiretrus anchorago (Fabricius) Collection dates: Townsend (1894) collected several nymphs (no adults) on 19 August 1891 identified by C.V. Riley. Counties: Doña Ana (?). Note: I have not seen specimens of this insect from New Mexico, though it also has been reported by Uhler (1876). Tylospilus acutissimus Stål Collection dates: 20 February through 8 October. Counties: Chaves, Colfax, Curry, Doña Ana, Eddy, Hidalgo, Lincoln, and Roosevelt. Note: A county record from Barber (1926) is included here. Zicrona americana Thomas Collection dates: 20 March through 26 August. Counties: Bernalillo, Chaves, Doña Ana, Lincoln, Otero, Roosevelt, Sandoval, Socorro, and Valencia. Note: Thomas (1992) described this species. Previous records of Z. caerulea (Linnaeus) in New Mexico likely all belong to Z. americana as Thomas (1992) examined individuals of this species from Arizona and Texas.

Subfamily Pentatominae Aelia americana Dallas Collection dates: 25 April through 13 August. Counties: Bernalillo, Colfax, Socorro, Torrance, and Valencia. Note: A county record from Ruckes (1937) is included here. Agonoscelis puberula Stål Collection dates: 28 May through 16 October. Counties: Catron, Doña Ana, Hidalgo, and Sierra. Note: This introduced African species first was reported from the United States, with records included from New Mexico, by Thomas et al. (2003). Antheminia remota (Horváth) Collection dates: 18 June through 6 July. Counties: Chaves, Colfax, Lincoln, and Taos. Note: Codophila remota (Horváth) was moved to Antheminia (see Rider 1998). County records from Barber (1926) and Ruckes (1937) are included here. Bagrada hilaris (Burmeister) Collection dates: 28 May through 16 October. Counties: Catron, Doña Ana, Otero, Socorro, and Valencia. Note: This introduced Old World species first was reported in California in 2008 (see Palumbo and Natwick 2010). It first was reported from New Mexico by Bundy et al. (2012). 200 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Banasa dimidiata (Say) Collection dates: 10 June through “August.” Counties: Santa Fe and Socorro. Note: A county record from Ruckes (1937) is included here. An alternate (original), but rarely used, spelling of this species (B. dimiata) was suggested by Froeschner (1988). However, Hoffman (2005) provides strong evidence for maintaining B. dimidiata as the correct spelling. Banasa euchlora Stål Collection dates: 27 March through 14 October. Counties: Bernalillo, Catron, Curry, Doña Ana, Eddy, Grant, Otero, Roosevelt, and Torrance. Banasa rolstoni Thomas and Yonke Collection dates: 14 April through 5 November. Counties: Doña Ana and Lincoln. Note: This is the first record of this insect from New Mexico. Banasa sordida (Uhler) Collection dates: 5 June through 8 September. Counties: Bernalillo, Catron, Doña Ana, Hidalgo, Santa Fe, and Taos. Note: A county record from Ruckes (1937) is included here. Banasa subcarnea Van Duzee Collection dates: 4 July through 5 November. Counties: Catron, Lincoln, Otero, and San Miguel. Note: A county record from Uhler (1904) is included here. Brochymena affinis Van Duzee Collection dates: 14 March through 23 November. Counties: Bernalillo, Doña Ana, Eddy, Lincoln, Otero, and Roosevelt. Note: This is the first report of this species from New Mexico. Brochymena dilata Ruckes Counties: Socorro. Note: I have not seen specimens of this insect from New Mexico, but it has been reported by Ruckes (1946). Brochymena hoppingi Van Duzee Collection dates: 20 March (1 ♀, this study). Counties: Doña Ana, Rio Arriba, and San Miguel. Note: County records from Ruckes (1937) are included here. Brochymena lineata Ruckes Collection dates: 27 March through 23 November. Counties: Catron, Doña Ana, McKinley, and Torrance. Note: A county record (and paratype) from Ruckes (1946) is included here. Brochymena myops Stål Collection dates: “July.” Counties: San Miguel. 2012 THE GREAT LAKES ENTOMOLOGIST 201

Note: I have not seen specimens of this insect from New Mexico. Ruckes (1946) accepted Uhler’s (1876) New Mexico record “with reservations,” possibly being a misidentified specimen of B. hoppingi. However, he did not mention his own identification of specimens collected previously from New Mexico (Ruckes 1937). Brochymena parva Ruckes Collection dates: 29 January through 21 December. Counties: Curry, Doña Ana, Los Alamos, Otero, and Sierra. Brochymena quadripustulata (Fabricius) Collection dates: 1 April through 18 November. Counties: Bernalillo, Chaves, Curry, Lea, Roosevelt, Santa Fe, Sierra, and Union. Note: County records from Ruckes (1937) are included here. Brochymena sulcata Van Duzee Collection dates: 10 February through 27 November. Counties: Bernalillo, Catron, Chaves, Curry, Doña Ana, Eddy, Lea, Luna, Otero, Quay, Sierra, and Valencia. Chinavia hilaris (Say) Collection dates: 29 February through 20 October. Counties: Bernalillo, Catron, Chaves, Doña Ana, Eddy, Otero, San Juan, Socorro, and Valencia. Note: The subgenus Chinavia within the genus Acrosternum was raised to genus (Schwertner 2005, Schwertner and Grazia 2006), transferring Acrosternum hilare (Say) to C. hilaris. This is the first record of this insect from New Mexico. Chlorochroa (Chlorochroa) congrua Uhler Collection dates: 8 May (2 adults) and 16 July (1 adult) 1891. Counties: Doña Ana. Note: The only records of this species from New Mexico are Townsend (1894), identified by Uhler. The presence ofC. congrua in the state needs to be verified. Chlorochroa (Chlorochroa) ligata (Say) Collection dates: 17 February through 10 December. Counties: Bernalillo, Chaves, Colfax, Curry, De Baca, Doña Ana, Eddy, Grant, Harding, Hidalgo, Lincoln, Otero, Roosevelt, Sierra, Socorro, Torrance, Union, and Valencia. Note: A county record from Barber (1926) is included here. Chlorochroa (Chlorochroa) opuntiae Esselbaugh Collection dates: 17 July through 12 October. Counties: Doña Ana, Eddy, and Roosevelt. Note: This is the first record of this insect from New Mexico. Chlorochroa (Chlorochroa) sayi (Stål) Collection dates: 10 February through 18 December. Counties: Bernalillo, Catron, Chaves, Colfax, Curry, De Baca, Doña Ana, Eddy, Guadalupe, Hidalgo, Lea, McKinley, Otero, Quay, Roosevelt, Socorro, Torrance, Union, and Valencia. 202 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Note: County records from Barber (1926) and Ruckes (1937) are included here. Chlorochroa (Chlorochroa) uhleri (Stål) Collection dates: 5 May 2001 (2 ♀♀, two counties, this study) and “June” (6 individuals) (Stroud 1950). Counties: Curry, Otero, and Roosevelt. Chlorochroa (Rhytidolomia) faceta (Say) Collection dates: 19 March through 24 November. Counties: Chaves, Colfax, Doña Ana, Hidalgo, Roosevelt, Socorro, and Valencia. Note: County records from Barber (1926) and Ruckes (1938) are included here. Chlorochroa (Rhytidolomia) osborni (Van Duzee) Collection dates: 13 January through 9 December. Counties: Chaves, Curry, Doña Ana, Eddy, Guadalupe, Roosevelt, Socorro, and Valencia. Chlorochroa (Rhytidolomia) viridicata (Walker) Collection dates: 27 May through 8 September. Counties: Chaves, Colfax, McKinley, Roosevelt, San Miguel, Santa Fe, and Socorro. Note: County records from Barber (1926) and Ruckes (1937) are included here. Chlorocoris (Chlorocoris) distinctus Signoret Note: I have not seen specimens of this insect from New Mexico, but it has been reported by Barber (1910), as C. atrispinus Stål. However, many specimens identified as C. distinctus actually are C. werneri Thomas (D. Thomas, pers. comm.). Chlorocoris (Monochrocerus) hebetatus Distant Collection dates: 9 June, 1992 (1 ♀). Counties: Hidalgo. Note: This is the first record of this insect from New Mexico. Coenus delius (Say) Collection dates: 4 May through 5 October. Counties: Bernalillo, Chaves, Curry, Doña Ana, Eddy, and Lincoln. Note: A county record from Rider (1995) is included here. Cosmopepla binotata Distant Collection dates: 8 June through 16 October. Counties: Bernalillo, Catron, Cibola, Doña Ana, Grant, Lincoln, Otero, Roosevelt, and Socorro. Note: A county record from Ruckes (1938) is included here. Cosmopepla decorata (Hahn) Collection dates: 16 April through 16 October. Counties: Catron, Doña Ana, Grant, Hidalgo, Lincoln, Otero, Roosevelt, Socorro, and Torrance. 2012 THE GREAT LAKES ENTOMOLOGIST 203

Note: McDonald (1986) first reported this species from New Mexico, no counties given. Cosmopepla lintneriana Kirkaldy Collection dates: “July and August” (no year given) Ruckes (1938), 10 August 1941 (2 ♂), and “September” 1959, (1 ♂) (current study). Counties: Doña Ana, Otero, and Santa Fe. Note: Though frequently in the literature as C. bimaculata (Thomas), its original combination is preoccupied; therefore, C. lintneriana is the correct name (Rider and Rolston 1995). Dendrocoris arizonensis Barber Note: Thomas and Brailovsky (1999) first reported this species from New Mexico, no counties given. Dendrocoris contaminatus Uhler Collection dates: 24 March through 3 November. Counties: Bernalillo, Chaves, Doña Ana, Lea, Lincoln, Luna, Otero, Santa Fe, and Sierra. Note: County records from Ruckes (1938) are included here. Dendrocoris humeralis (Uhler) Collection dates: 6 August (no year given, 4 specimens). Counties: San Miguel. Note: The only record of this species from New Mexico is Uhler (1904). The presence of D. humeralis in the state needs to be verified. Dendrocoris nelsoni Thomas and Brailovsky Collection dates: 18 May 1985 (1 ♀). Counties: Lincoln. Note: A single specimen (paratype) has been reported from New Mexico by Thomas and Brailovsky (1999). Dendrocoris neomexicanus Nelson Collection dates: 8 June through 18 August. Counties: McKinley and Otero. Note: County records (and types) from Nelson (1955) are included here. Dendrocoris parapini Nelson Collection dates: 1 July through 16 August. Counties: San Miguel, Sandoval, and Santa Fe. Note: County records (and types) from Nelson (1957) are included here. Dendrocoris pini Montandon Note: I have not seen specimens of this insect from New Mexico, but it has been reported by Nelson (1955), no counties given. Dendrocoris reticulatus Barber Collection dates: 13 (no year given, 2 specimens) (Uhler 1904) and 18 August 1993 (2 ♀♀, this study). Counties: Hidalgo and San Miguel. Euschistus biformis Stål Collection dates: 20 April through 4 December. 204 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Counties: Doña Ana, Grant, Hidalgo, and Otero. Note: This is the first record of this species from New Mexico. Euschistus egglestoni Rolston Collection dates: 5 August through 30 October. Counties: Doña Ana, Hidalgo, and Lea. Note: This is the first record of this species from New Mexico. Euschistus inflatus Van Duzee Collection dates: 8 March through 19 October. Counties: Catron, Doña Ana, Grant, Lincoln, Roosevelt, San Miguel, Sierra, and Valencia. Note: A county record each from Ruckes (1937) and Van Duzee (1903) are included here. Euschistus servus servus (Say) Collection dates: 2 April through 28 October. Counties: Catron, Curry, Doña Ana, Eddy, Grant, Hidalgo, Luna, Otero, Roosevelt, and Valencia. Note: County records from Ruckes (1938) are included here. Euschistus variolarius (Palisot) Counties: Bernalillo. Note: I have not seen specimens of this insect from New Mexico, but it has been reported by Osborn (1893) from a single locality. The presence of E. variolarius in the state needs to be verified. Halyomorpha halys (Stål) Collection dates: 7 November 2011 (2 ♀♀). Counties: Luna. Note: These two specimens of the brown marmorated stink bug were collected on a trailer from Pittsburg, Pa. This is the first record of this species from New Mexico. Holcostethus abbreviatus Uhler Collection dates: 18 April through 27 July. Counties: Doña Ana, Grant, Santa Fe, and Union. Note: A county record from Ruckes (1938) is included here. Holcostethus limbolarius (Stål) Collection dates: 7 March through 29 October. Counties: Cibola, Colfax, Doña Ana, Eddy, Grant, Hidalgo, Lea, Lin- coln, Otero, Roosevelt, San Miguel, Santa Fe, Sierra, and Socorro. Note: County records from Ruckes (1937) and Uhler (1904) are included here. Holcostethus ruckesi McDonald Collection dates: 18 July 1961 (1 ♀). Counties: Grant. Note: A single specimen (paratype) has been reported from New Mexico by McDonald (1974). 2012 THE GREAT LAKES ENTOMOLOGIST 205 Hymenarcys crassa Uhler Collection dates: 8 April 1993 (1 ♀). Counties: Hidalgo. Note: This is the first record of this species from New Mexico. Loxa flavicollis (Drury) Note: Uhler (1876) reported seeing specimens collected in New Mexico. The presence of L. flavicollisin the state needs to be verified. Mecidea major Sailer Collection dates: The insect has been collected every month of the year. Counties: Bernalillo, Chaves, Doña Ana, Grant, Hidalgo, Otero, Quay, Roosevelt, Sierra, Socorro, Union, and Valencia. Mecidea minor Ruckes Collection dates: This insect has been collected every month of the year. Counties: Bernalillo, Chaves, Doña Ana, Grant, Hidalgo, Otero, Quay, Roosevelt, Sierra, and Valencia. Menecles insertus (Say) Collection dates: 7 March through 20 October. Counties: Cibola, Bernalillo, Doña Ana, Eddy, Otero, Roosevelt, and Sierra. Note: This is the first record of this species from New Mexico. Mcphersonarcys aequalis (Say) Collection dates: 14 August 1962 (1 ♀). Counties: Hidalgo. Note: Hymenarcys aequalis was placed in a new genus by Thomas (2012). This is the first record of this species from New Mexico. Mormidea cubrosa Dallas Note: I have not seen specimens of this insect from New Mexico, but it has been reported by Uhler (1876). Moromorpha tetra (Walker) Collection dates: 11 June through 30 October. Counties: Doña Ana, Eddy, Grant, Hidalgo, Lea, Lincoln, and Luna. Murgantia histrionica (Hahn) Collection dates: 28 February through 22 November. Counties: Bernalillo, Catron, Chaves, Colfax, De Baca, Doña Ana, Eddy, Grant, Guadalupe, Harding, Luna, Otero, Roosevelt, Sandoval, San Miguel, Sierra, Torrance, and Valencia. Note: A county record from Barber (1926) is included here. Neottiglossa sulcifrons Stål Collection dates: “15-30 July” (no year given) (Ruckes 1938). Counties: Doña Ana, Hidalgo, and Luna. Nezara viridula (Linnaeus) Collection dates: 21 September 1971 (1 ♂). 206 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Counties: Roosevelt. Note: This is the first record of this species from New Mexico. Oebalus mexicanus (Sailer) Collection dates: 27 July through 5 November. Counties: Doña Ana and Grant. Note: This is the first record of this insect from New Mexico. Oebalus pugnax (Fabricius) Collection dates: This insect has been collected from 5 June through “October.” Counties: Bernalillo and Roosevelt. Parabrochymena arborea (Say) Collection dates: “June” through 1 August. Counties: Hidalgo, San Miguel, and Santa Fe. Note: County records from Ruckes (1937) are included here. Larivière (1992) erected the genus, and Ahmad and McPherson (1998) supported the validity of Parabrochymena. Piezodorus guildinii (Westwood) Note: Van Duzee (1904) reported seeing an individual collected from New Mexico. The presence of P. guildinii in the state needs to be verified. Prionosoma podopioides Uhler Collection dates: 3 April through 23 October. Counties: Bernalillo, Chaves, Colfax, Doña Ana, Eddy, Otero, Roos- evelt, and Socorro. Note: County records from Ruckes (1937) and Stroud (1950) are included here. Tepa brevis (Van Duzee) Collection dates: 7 June through 13 October. Counties: Doña Ana, Otero, and Roosevelt. Note: A county record from Stroud (1950) is included here. Tepa rugulosa (Say) Collection dates: 29 April through 20 October. Counties: Chaves, Colfax, Roosevelt, and San Juan. Note: A county record from Ruckes (1937) is included here. Tepa vanduzeei Rider Collection dates: 18 June through 6 August. Counties: Sandoval, San Juan and Taos. Note: A county record from Rider (1986) is included here. Tepa yerma (Rolston) Collection dates: 10 April through 20 October. Counties: Cibola, Doña Ana, and Socorro. Note: This is the first record of this species from New Mexico. 2012 THE GREAT LAKES ENTOMOLOGIST 207 Thyanta custator accerra McAtee Collection dates: 3 March through 23 November. Counties: Bernalillo, Catron, Chaves, Cibola, Colfax, Curry, De Baca, Doña Ana, Eddy, Grant, Guadalupe, Harding, Hidalgo, Lea, Lincoln, Luna, McKinley, Otero, Quay, Rio Arriba, Roosevelt, Sandoval, San Miguel, Santa Fe, Socorro, Torrance, Union, and Valencia. Note: County records from Rider and Chapin (1992) are included here. Trichopepla semivittata (Say) Collection dates: 8 August through 12 October. Counties: Chaves, Doña Ana, and Roosevelt. Note: This is the first record of this species from New Mexico.

Acknowledgments I thank Sandra Brantley (Museum of Southwestern Biology, Division of Arthropods, University of New Mexico) and Darren Pollock (Natural History Museum, Eastern New Mexico University) for providing access to material from their respective collections, and Daniel Swanson (Museum of Zoology, University of Michigan) for access to NM records housed in the collections at Michigan State and the University of Michigan. I thank Don Thomas (USDA ARS) for review of the manuscript and identifications of some specimens, and Max Bleiweiss (NMSU) for generating the state map. This research was partially funded by the New Mexico State Agricultural Experiment Station, Las Cruces.

Literature Cited Ahmad, I., and J. E. McPherson. 1998. Additional information on male and female genitalia of Parabrochymena Larivière and Brochymena Amyot & Serville (Hemip- tera: Pentatomidae). Annals of the Entomological Society of America 91: 800–807. Barber, G. W. 1926. A short list of Scutelleroidea collected in New Mexico in 1916. Entomological News 37: 43–44. Barber, H. G. 1910. Some Mexican Hemiptera-Heteroptera new to the fauna of the United States. Journal of the New York Entomological Society 18: 34–39. Bundy, C. S., T. R. Grasswitz, and C. Sutherland. 2012. First report of the invasive stink bug Bagrada hilaris (Burmeister) (Heteroptera: Pentatomidae) from New Mexico, with notes on its biology. Southwestern Entomologist 37: 411–414. Froeschner, R. C. 1988. Family Pentatomidae Leach, 1815. The stink bugs, pp. 544–597. In T. J. Henry, and R. C. Froeschner (eds.), Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, New York. 958 pp. Hoffman, R. L. 2005. The Virginia species of Banasa, three decades later (Heteroptera: Pentatomidae). Banisteria 25: 41–44. Larivière, M.-C. 1992. Description of Parabrochymena, new genus, and redefinition and review of Brochymena Amyot and Audinet-Serville (Hemiptera: Pentatomidae), with considerations on natural history, chorological affinities, and evolutionary relationships. Memoirs of the Entomological Society of Canada 163: 1–75. McDonald, F. J. D. 1974. Revision of the genus Holcostethus in North America (Hemip- tera: Pentatomidae). Journal of the New York Entomological Society 82: 245–258. McDonald, F. J. D. 1986. Revision of Cosmopepla Stål (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 94: 1–15. 208 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Nelson, G. H. 1955. A revision of the genus Dendrocoris and its generic relationships (Hemiptera: Pentatomidae). Proceedings of the Entomological Society of Washington 57: 49–67. Nelson, G. H. 1957. A new species of Dendrocoris and a new combination of Atizies (He- miptera: Pentatomidae). Proceedings of the Entomological Society of Washington 59: 197–199. Osborn, H. 1893. Notes on the distribution of Hemiptera. Proceedings of the Iowa Acad- emy of Sciences 1 (part 3): 120–123. Palumbo, J. C., and E. T. Natwick. 2010. The Bagrada bug (Hemiptera: Pentatomidae): A new invasive pest of cole crops in Arizona and California. Online. Plant Health Progress doi:10.1094/PHP-2010-0621-01-BR. Rider, D. A. 1986. A new species and new synonymy in the genus Tepa Rolston and McDonald (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 94: 552–558. Rider, D. A. 1995. Review of the genus Coenus Dallas, with the description of C. explanatus, new species (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 103: 39–47. Rider, D. A. 1998. Nomenclatural changes in the Pentatomoidea (Hemiptera-Heteroptera: Cydnidae, Pentatomidae). II. Species level changes. Proceedings of the Entomological Society of Washington 100: 449–457. Rider, D. A. 2009. Pentatomoidea Home page. http://www.ndsu.nodak.edu/ndsu/rider/ Pentatomoidea; accessed 3 July, 2012. Rider, D. A., and J. B. Chapin. 1992. Revision of the genus Thyanta Stål, 1862 (Het- eroptera: Pentatomidae) II. North America, Central America, and the West Indies. Journal of the New York Entomological Society 100: 42–98. Rider, D. A., and L. H. Rolston. 1995. Nomenclatural changes in the Pentatomidae (Hemiptera-Heteroptera). Proceedings of the Entomological Society of Washington 97: 845–855. Ruckes, H. 1937. An annotated list of some pentatomids (Heteroptera) from New Mexico. Bulletin of the Brooklyn Entomological Society 32: 32–36. Ruckes, H. 1938. Additions to an annotated list of pentatomids (Heteroptera) of New Mexico. Bulletin of the Brooklyn Entomological Society 33: 10–13. Ruckes, H. 1946. Notes and keys on the genus Brochymena (Pentatomidae, Heteroptera). Entomologica Americana 26: 143–238. Schwertner, C. F. 2005. Filogenia e classificação dos percevejos-verdes do grupoNezara Amyot & Serville 1843 (Hemiptera, Pentatomidae, Pentatominae). Ph.D. dissertation, Universidade Federal do Rio Grande do Sul, Porto Alegre. xv+246 pp. Schwertner, C. F. and J. Grazia. 2006. Descrição de seis espécies de Chinavia (He- miptera, Pentatomidae, Pentatominae) da América do Sul. Iheringia Serie Zoologia, Porto Alegre 96: 237–248. Stroud, C. P. 1950. A survey of the insects of White Sands National Monument, Tularosa Basin, New Mexico. American Midland Naturalist 44: 659–677. Thomas, D. B. 1992. Taxonomic synopsis of the Asopinae Pentatomidae (Heteroptera) of the Western Hemisphere. Thomas Say Foundation Monographs, Vol. 15. Entomologi- cal Society of America, Lanham. 156 pp. Thomas, D. B. 2012. Mcphersonarcys, a new genus for Pentatoma aequalis Say (Heter- optera: Pentatomidae). The Great Lakes Entomologist 45: 127–133. Thomas, D. B., and H. Brailovsky. 1999. Review of the genus Dendrocoris Bergroth with descriptions of new species (Pentatomidae: Heteroptera). Insecta Mundi 13: 1–9. 2012 THE GREAT LAKES ENTOMOLOGIST 209

Thomas, D. B., J. E. Eger, W. Jones, and G. Ortega-Leon. 2003. The African cluster bug, Agonoscelis puberula (Heteroptera: Pentatomidae), established in the New World. Florida Entomologist 86: 151–153. Townsend, C. H. T. 1894. Notes on some south-western Hemiptera. The Canadian Entomologist 26: 312–316. Uhler, P. R. 1871. A list of Hemiptera collected in eastern Colorado,and Northeastern New Mexico, by C. Thomas, during the expedition of 1869, pp. 471–472. In Hayden, F. V. Preliminary Report of the United States Geological Survey of Wyoming and Portions of Contiguous Territories, Second Annual Progress Report. Uhler, P. R. 1876. List of Hemiptera of the region west of the Mississippi River, including those collected during the Hayden explorations of 1873. Bulletin of the United States Geological and Geographical Survey of the Territories 1: 267–361, plates 1–21. Uhler, P. R. 1904. List of Hemiptera-Heteroptera of Las Vegas Hot Springs, New Mexico, collected by Messrs. E. A. Schwarz and Herbert S. Barber. Proceedings of the United States National Museum 27: 349–364. Van Duzee, E. P. 1903. Hemiptera of Beulah, New Mexico. Transactions of the American Entomological Society 29: 107–113. Van Duzee, E. P. 1904. Annotated checklist of the Pentatomidae recorded from America north of Mexico, with descriptions of some new species. Transactions of the American Entomological Society 30: 1–80. 210 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 The Pentatomidae, or Stink Bugs, of Kansas with a key to species (Hemiptera: Heteroptera) Richard J. Packauskas1

Abstract Forty eight species of Pentatomidae are listed as occurring in the state of Kansas, nine of these are new state records. A key to all species known from the state of Kansas is given, along with some notes on new state records.

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The family Pentatomidae, comprised of mainly phytophagous and a few predaceous species, is one of the largest families of Heteroptera. Some of the phytophagous species have a wide host range and this ability may make them the most economically important family among the Heteroptera (Panizzi et al. 2000). As a group, they have been found feeding on cotton, nuts, fruits, vegetables, legumes, and grain crops (McPherson 1982, McPherson and McPherson 2000, Panizzi et al 2000). On soybeans alone, losses can accrue to $60 million in the southeastern United States (McPherson and McPherson 2000), while in Georgia, in 1985, pecan industry losses to stink bugs were estimated to run as high as $3.5 million (Douce and Suber1986). A recent introduction (first seen in 1996 in PA), the brown marmorated stinkbug (BMSB, Halyomorpha halys (Stål)), has been found invading homes as overwintering sites in record numbers in the east and Pacific coast and becoming a major pest on a wide variety of crops, fruit trees, and ornamentals (Hamilton 2009; Nielsen and Hamilton 2009a, b; USDA NAL 2012). It has not been found in Kansas yet, but has been detected in 36 states and continues to expand its range. It is a striking brown colored species with alternating white and black bands on its antennae as well as similar markings along the sides of the abdomen. The majority of the literature on pentatomids has been in reference to the eastern portion of the United States, with few publications on the family from the western states. This paper catalogs the species of Kansas and a key is given for identification of those species. Prior to this study, 35 species of pentatomids were listed from Kansas by Froeschner (1988). Here I expand that number to 48, and include notes on 9 new state records. I have not included Popenoe’s (1885) record of Chinavia pennsylvanica (Gmelin) (as Rhaphigaster pennsylvanicus), which would have been the furthest western record of this species, nor Crevecoeur’s (1905) record of Perillus confluens (Herrich-Schaeffer); both records are not included in Froeschner’s (1988) localities and reference locality and specimen data is lacking for both.

Materials and Methods The body of this work comes from examination and identification of specimens in both the Fort Hays Sternberg Museum (at Fort Hays State University) and the University of Kansas Biodiversity Institute Entomological Collection. For distributions of species, I relied mainly on Froeschner (1988), but supplemented

1Department of Biological Sciences, Fort Hays State University, 600 Park St. Hays, KS 67601. 2012 THE GREAT LAKES ENTOMOLOGIST 211 this with other works that also facilitated identification (Knight, 1952; Lariviére, 1992; McDonald,1974, 1976, 1986; McPherson, 1982; McPherson and McPherson, 2000; Rider 1986, 1995; Rider and Chapin, 1992; Rolston and McDonald, 1984; Thomas, 1983, 1992; Thomas and Yonke 1981; Zimmer1910, 1912). Key. The following key to species of Kansas was mainly modified from McPherson (1982), but also aided by McPherson and McPherson (2000), Bux- ton et al. (1983), Larivière (1992), McDonald (1986), Rider (1986, 1995), Rider and Chapin (1992), Rolston (1972, 1974), Rolston and McDonald (1984), Sailer (1952), Thomas (1983, 1992), and Zimmer (1910, 1912). Recent nomenclatorial changes include the creation of a new genus, McPhersonarcys Thomas, for Hy- menarcys aequalis (Say) by Thomas (2012, this issue) and the recent placement of Acrosternum hilare (Say) into the genus Chinavia Orian (see Schwertner and Grazia 2006). A list of all species can be seen in Table 1. Key to the species of Pentatomidae from Kansas 1 Eyes stalked (pedunculate); scutellum broadly U-shaped, nearly reaching tip of abdomen; lacking bright, contrasting colors (Podopinae, Amaurochrous)...... 2 1′ Eyes not stalked; scutellum usually roundedly triangular, but if large and U-shaped then having a broad oval shape (Coenus spp.) or bright, contrasting colors...... 3 2(1) Juga and tylus equal or subequal at apices; lateral margin of pronotum strongly sinuate...... Amaurochrous cinctipes (Say)

Table 1. Pentatomidae of Kansas

* = new state record (also bolded)

Subfamily Asopinae Euschistus latimarginatus Zimmer* Alcaeorrhynchus grandis (Dallas)* Euschistus servus servus (Say) Apoecilus cynicus (Say) Euschistus tristigmus tristigmus (Say) Perillus bioculatus (Fabricius) Euschistus variolarius (Palisot) Podisus maculiventris (Say) Holcostethus abbreviatus Uhler Stiretrus anchorago (Fabricius) Holcostethus limbolarius (Stål) Zicrona caerulea (Linnaeus)* Hymenarcys nervosa (Say)* Mcphersonarcys aequalis (Say) SubfamilyPentatominae Mecidea major Sailer Acrosternum hilare (Say) Menecles insertus (Say) Aelia americana Dallas Mormidea lugens (Fabricius) Banasa dimidiata (Say)* Murgantia histrionica (Hahn) Banasa euchlora Stål * Neottiglossa sulcifrons Stål Brochymena cariosa Stål Oebalus pugnax pugnax (Fabricius) Brochymena quadripustulata (Fabricius) Parabrochymena arborea (Say) Brochymena sulcata Van Duzee* Prionosoma podopioides Uhler * Chinavia hilare (Say) Tepa vanduzeei Rider Chlorochroa faceta (Say) Thyanta custator acerra McAtee Chlorochroa ligata (Say) Thyanta pallidovirens (Stål)* Chlorochroa persimilis Horvath Trichopepla atricornis Stål* Chlorochloa sayi (Stål)* Trichopepla semivittata (Say) Chlorochroa uhleri (Stål) Tylospilus acutissimus (Stål)* Coenus delius (Say) Cosmopepla intergressus (Uhler) Subfamily Podopinae Cosmopepla lintneriana (Thomas) Amaurochrous brevitylus Barber & Sailer Dendrocoris humeralis (Uhler) Amaurochrous cinctipes (Say) 212 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

2′ .Juga markedly longer than tylus, lateral margin of pronotum nearly straight...... Amaurochrous brevitylus Barber & Sailer 3(1) First segment of beak short, thickened, often free, extending well beyond bucculae; bucculae converging beneath basal segment of beak (Asopinae)...... 4 3′ First segment of beak slender, lying between bucculae (occasionally extending slightly beyond bucculae); bucculae subparallel, not converging posteriorly beneath basal segment (Pentatominae)...... 10 4(3) Humeri extended as a bifid process, length over 15 mm...... Alcaeorrhynchus grandis (Dallas) 4′ Humeri acute, angled, or rounded, never extended as a bifid process, length less than 10 mm...... 5 5(4) Profemora armed with ventral spine or tubercle at distal third to fourth; males with submedian pubescent patch on abdominal sternites 4-6...... 6 5′ Profemora unarmed; males lacking submedian pubescent patches on abdominal sternites...... 7 6(5) Scutellum U-shaped, broadly rounded apically and its tip nearly reaching apex of abdomen; frena about ¼ length of scutellum...... Stiretrus anchorago (Fabricius) 6′ Scutellum not U-shaped, its tip not reaching apex of abdomen; frena about ½ length of scutellum...... Perillus bioculatus (Fabricius) 7(5) Abdominal sternite 2 (first visible) unarmed; color metallic purplish- blue to black; male pygophore lacking brushlike setae, at either side of middle on ventroposterior rim...... Zicrona americana Thomas 7′ Abdominal sternite 2 armed with slender anteriorly directed spine; color not metallic blue to black...... 8 8(7) Juga slightly exceeding tylus in length; overall length usually more than 14mm...... Apoecilus cynicus (Say) 8′ Juga equal in length to tylus; overall length less than 12mm...... 9 9(8) Scutellum with 3 whitish maculae at base; black macula centrally on each corium and median black stripe extending from tip of scutellum to apex of membrane...... Tylospilus acutissimus (Stål) 9′ Without above combination of colors, dull brown with apex of corium slightly to fully red, often extending along margin...... Podisus maculiventris (Say) 10(3) Body length to width ratio 3:1 or more; abdominal venter with wide longitudinal band of striations (stridulatory patch, often difficult to view) on first 3 segments on either side of middle (Mecideini)Mecidea ...... 46 10′ Body length to width ration 2.5:1, or much less; abdominal venter without striated areas...... 11 11(10) Juga each with subapical, lateral tooth; pronotum strongly dentate anterolaterally; abdominal venter with median shallow, longitudinal depression, disappearing posteriorly (Halyini)...... 12 11′ Juga lacking subapical teeth; pronotum variable anterolaterally, rarely dentate; abdomen without median ventral depression (at most, on basal segment) (Pentatomini)...... 14 1 12(11) Scutellar base (¼ to /3) elevated above remainder; humeri quadrate with large teeth...... Parabrochymena arborea (Say) 1 12′ Scutellar base (¼ to /3) slightly elevated; humeri subtriangular with small teeth (Brochymena)...... 13 2012 THE GREAT LAKES ENTOMOLOGIST 213 13(12) Juga distinctly longer than tylus, exceeding it by at least their width at that point...... Brochymena quadripustulata (Fabricius) 13′ Juga subequal to tylus, if exceeding it then by less than their width at that point...... Brochymena cariosa Stål 14(11) First visible segment of abdomen with a median spine or slightly raised area anteriorly directed (except in males of Dendrocoris humeralis).... 15 14′ First visible segment of abdomen without anteriorly directed spine or raised area...... 18 15(14) Juga extended and usually touching beyond tylus...... Dendrocoris humeralis (Uhler) 15′ Juga not extending beyond tylus, but if so, then not touching...... 16 16(15) Large, over 13 mm; green; first visible abdominal segment with anteriorly directed acute spine...... Chinavia hilare (Say) 16′ Smaller, less than 10 mm; green or brown; first visible abdominal segment with a median raised area or obtuse spine directed anteriorly (Banasa)...... 17 17(16) Green in color; pronotum without distinctly darker area posteriorly; scutellum with whitish spot at each angle...... Banasa euchlora Stål 17′ Brown to green in color; pronotum lighter anteriorly, with a distinctly darker area beyond a line drawn through humeri, scutellum without light spots...... Banasa dimidiata (Say) 18(14) Ostiole usually with distinct auricle (sometimes not evident), but not extending as a long tapering canal, ostiole acutely tapering on inner side...... 19 18′ Ostiole without auricle, but extending laterally as a long, tapering canal, ostiole rounded on inner side…...... 35 19(18) Ostiole small (slight depression), without auricle; juga with whitish spot on either side of tylus; color variegated black with red to yellow...... Murgantia histrionica (Hahn) 19′ Ostiole conspicuous, auricle usually well-developed (lacking or small in Neottiglossa), color not as above…...... 20 20(19) Posterior margin of humeral angle emarginate; body heavily setose...... Prionosoma podopioides Uhler 20′ Posterior margin of humeral angle not emarginated; body not heavily setose...... 21 21(20) Scutellum shorter than corium, tip not broadly rounded, with apical third narrower than apex of corium….…...... 22 21′ Scutellum longer or equal to corium, tip broadly rounded, with apical third wider than apex of corium….…...... 30 22(21) Hind tibia with distinct sulcus dorsally for entire length...... 23 22′ Hind tibia without distinct sulcus for entire length...... 29 23(22) Pronotal margin anterolaterally crenulate; humeral angle acute to rounded (Euschistus)...... 24 23′ Pronotal margin anterolaterally not crenulate, humeral angles broadly rounded…...... 27 24(23) Abdominal sternites with black maculae at lateral angles; humeri variable; antennae with segments 4 and 5 pale red, or brown to black; males without dark macula on ventral side of pygophore...... 26 214 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

24′ Abdominal sternites without black maculae at lateral angles; humeri subacute to spinose; antennae with apical half of segment 4 and all of 5 black...... 25 25(24) Juga longer than tylus, apex of head incised; membrane immaculate (from original description).….Euschistus latimarginatus Zimmer 25′ Juga subequal to tylus; membrane dotted with small brown maculae; males with dark brown to black macula on ventral side of pygophore...... Euschistus variolarius (Palisot de Beauvois) 26(24) Abdominal venter with 1 to 4 median black maculae, occasionally obsolete; length less than 12 mm.....Euschistus tristigmus tristigmus (Say) 26′ Abdominal venter without black maculae; length variable...... Euschistus servus servus (Say) 27(23) Pronotum with anterior angles strongly produced, broadly widened, reaching level of mideye, margins strongly explanate, head appears recessed into prothorax; length 12 or more mm...... Menecles insertus (Say) 27′ Pronotum with anterior angles less widened, never reaching level of mideye, margins not strongly explanate; length 11.5 mm or less.....28 28(27) Head with sides parallel in front of middle, its length subequal to width across eyes; body length 6.0-8.5 mm...... Mcphersonarcys aequalis (Say) 28′ Head with sides sinuos in front of middle, its length much shorter than width across eyes; body length 8.5-11.5 mm....Hymenarcys nervosa (Say) 29(22) Pronotal humeri spined, directed anteriorly, beak segment 1 not longer than bucculae...... Oebalus pugnax (Fabricius) 29′ Pronotal humeri not spined; beak segment 1 distinctly longer than bucculae ...... Mormidea lugens (Fabricius) 30(21) Tylus distinctly elevated above juga; membrane of hemelytron pale with strongly anastomosing veins (Coenus)...... 31 30′ Tylus scarcely, if at all, elevated above juga; membrane without strongly anastomosing veins...... 32 31(30) Brown spots on upper surface of tibiae small, limited to base of each seta; posterior margin of male pygophore with distinct medial tooth...... Coenus delius (Say) 31′ Fuscous spots on upper surface of tibiae large, irregular; posterior margin of male pygophore lacking medial tooth...... Coenus inermis Harris & Johnston 32(30) Juga subequal to tylus: body black to brown with red or orange markings, head black...... 33 32′ Juga longer than tylus, body without red or orange markings, head not black...... 34 33(32) Black with red markings, scutellum with two distinct marginal red spots...... Cosmopepla lintneriana (Kirkaldy) 33′ Brown or reddish brown with markings in orange and black; scutellum lacking red spots...... Cosmopepla intergressa (Uhler) 34(32) Propleura expanded anteriorly as thin plate surpassing level of base of antenna; longitudinal yellow levigate line extending from head to scutellum, bordered by black bands...... Aelia americana Fabricius 34′ Propleura not extended anteriorly as a thin plate, not surpassing base of eye; two prominent levigate yellow spots at basal angles of scutellum...... Neottiglossa sulcifrons Stål 2012 THE GREAT LAKES ENTOMOLOGIST 215 35(18) Juga longer than tylus by distance longer than width of tylar apex (Holcostethus)...... 36 35′ Juga slightly shorter to slightly longer than tylus, if longer then exceeding tylus by less than width of tylar apex...... 37 36(35) Connexiva black with a pale margin; scutellum narrowed at tip; antennal segments 1-3 yellow to red, 4 and 5 brown to blackish...... Holcostethus limbolarius (Stål) 36′ Connexiva alternated black and yellow; scutellum broader at apex; antennal segments concolorous...... Holcostethus abbreviatus Uhler 37(35) Body distinctly pubescent, especially sides of abdominal sternites; frena reaching to middle of scutellum (Trichopepla)...... 38 37′ Body not pubescent, or slightly so; frena surpassing middle of scutellum...... 39 38(37) Head narrowly rounded, tapering apically; antennae reddish brown, 2 distal segments black...... Trichopepla semivittata (Say) 38′ Head broadly rounded or truncate apically, nearly porrect; antennae black with only basal segment pale...... Trichopepla atricornis Stål 39(37) Bucculae lobed or squarely truncate posteriorly; length of body 7.5 mm or less...... Tepa vanduzeei Rider 39′ Bucculae gradually disappearing or arcuate posteriorly; length, 10 mm or more...... 40 40(39) Ostiolar ruga reaching more than ½ distance from inner margin of ostiole to lateral margin of metapleuron (Thyanta)...... 41 40′ Ostiolar ruga reaching ½ or less distance from inner margin of ostiole to lateral margin of metapleuron (Chlorochroa)...... 42 41(40) Anterolateral margins and mesial angle of cicatrices of pronotum black...... Thyanta calceata (Say) 41′ Anterolateral margins of pronotum not black, color of mesial angle of cicatrices variable, often concolorous to pronotum...... Thyanta custator accerra McAtee 42(40) Scutellum with yellow medial stripe (rarely faint)...... Chlorochloa faceta (Say) 42′ Scutellum without yellow medial stripe...... 43 43(42) Scutellum with 3 ivory colored callosities at base; embolium subparallel or slightly widened apically, apex less than twice as wide as base...... 44 43′ Scutellum lacking 3 ivory colored callosities at base, tip of scutellum whitish to red; embolium widened apically, with apex nearly twice as wide as base...... 45 44(43) Embolium wider apically; hemelytral membrane with purple flecks; callosities at base of scutellum moderate to small...... Chlorochloa uhleri (Stål) 44′ Embolium parallel, not widened apically; hemelytral membrane lacking purple flecks; callosities at base of scutellum large, distinct...... Chlorochloa sayi (Stål) 45(43) Dark tan to nearly black species, with margins of both pronotum and abdomen red to yellow...... Chlorochloa ligata (Say) 45′ Green colored species, with margins whitish yellow to red...... Chlorochloa persimilis Horvath 216 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

46(10) Males with medial tubercle slightly ventrad of ventroposterior rim of pygophore; female abdominal lateral black spots as wide as one-twelfth width of supporting segment...... Mecidea minor Ruckes 46′ Males lacking medial tubercle slightly ventrad of ventroposterior rim of pygophore; female abdominal lateral black spots as wide as one-eighth width of supporting segment...... Mecidea major Sailer

Notes on New State Records (NSR) and other species notes The new state records were all determined by Packauskas, unless otherwise noted and are housed at the Fort Hays Sternberg Museum (FHSM), Fort Hays State University, University of Kansas Biodiversity Institute (KUBI), entomology collection, or University of Michigan Museum of Zoology (UMMZ). Alcaeorrhynchus grandis (Dallas), NSR. Published records for this species are restricted to FL, AR, and TX, although Barton and Lee (1981) suggested the Arkansas specimens might have been accidental introductions. Six specimens (FHSM): 1999-2007, Ellis County. Banasa dimidiata (Say), NSR. This species may occur across the entire U. S. continent, and has been recorded previously in NE and OK. Froeschner (1988) noted that the proper specific epithet should bedimiata ; however, Hoff- man (2005) provided strong evidence for accepting the emendation to dimidiata and I follow McPherson and McPherson (2000) in calling it B. dimidiata. Four specimens (FHSM): 1964-2005, Cherokee and Ellis County. Banasa euchlora Stål, NSR. This species was previously known from CO, NV, UT, TX and OK, as well as numerous midwestern states. Eleven specimens (FHSM): 1995-2008, Barton, Ellis, Harper, McPherson, Rooks, and Russell County. Brochymena carolinensis (Westwood). Although Lariviére (1992) published a record for Kansas as “Eagleton,” this should be attributed to Eagleton, OK, as there is no Eagleton, KS. Lariviére also refers to OK for this species, although this is not listed in her distribution list. Chlorochloa sayi (Stål), NSR. This species is mainly western in distribution, but has been recorded from CO and AR. Three specimens (FHSM), 16 (KUBI): 1902-1998, Hamilton, Morton, and Trego County. Coenus inermis Harris and Johnson. Rider (1996) reported a single specimen from Montgomery County. Euschistus latimarginatus Zimmer, NSR. This species has only been recorded from CO and NE. Four specimens (KUBI, determined by D. Stoner): 1902-1912, Clark, Rooks, Russell, and Thomas County. Hymenarcys nervosa (Say). This species is mostly eastern in distribution, but has been recorded in OK and TX, while Crevecoeur (1905) recorded this from KS. This is now confirmed by a single specimen (UMMZ, det. R. F. Hussey, 1952): 1927, Franklin County. (personal communication, Daniel Swanson). Mecidea minor Ruckes. Sailer (1952) reported specimens from Clark, Hamilton, Meade, Morton, Saint John, Scott, Seward, and Stevens County, but these KS records were apparently overlooked by Froeschner (1988). Prionosoma podopioides Uhler, NSR. This species was recorded from NE and CO, and is known from other western states as well as IA, IL, and MI. Twelve specimens (KUBI, det. J. D. Lattin, 1958): 1911-1952, Douglas, Ells- worth, Kiowa, Meade, Norton, Reno, and Stafford County. Thyanta calceata (Say). Rider and Chapin (1992) reported specimens from Bourbon and Douglas County. 2012 THE GREAT LAKES ENTOMOLOGIST 217

Trichopepla atricornis Stål, NSR. This species seems to be northwestern in distribution, but has been recorded from CO, UT, and WY. It is easily separated from T. semivittata in having 4 longitudinal rows of black maculae across abdominal sterna and 4 ultimate segments of the antennae black, as well as the connexival segments with a pale margin (not variegated, occasionally uniformly brown). Three specimens (FHSM): 2000-2004, Ellis County. Tylospilus acutissimus (Stål), NSR. This species is only known from AZ, CO, NM, and TX. One specimen, lacking a head and pronotum, but nonetheless distinct (KUBI, det. by K. A. Philips, 1982): 1944, Ford County. Zicrona americana Thomas, NSR. This species has been recorded by Thomas (1992) from AZ, CA, and TX, and is relatively uncommon. One specimen: 2007, Logan County.

Acknowledgments This paper was written in honor of Dr. J. E. McPherson, who is retiring from his position at Southern Illinois University at Carbondale, Illinois. He was my Master’s degree advisor and put me on track to study Heteroptera. Jay has finally gotten his way in having many of his former students all work on pentatomids. I would like to give my thanks to Zach Falin, collections man- ager, for allowing me some time in the collections of the University of Kansas Biodiversity Institute. Dr. Dave Rider deserves a special thank you for criti- cally reviewing this paper, forcing it to be better. I would also like to sincerely thank Sheran Powers, Interlibrary Loan Coordinator, at the Forsyth Library, Fort Hays State University, for her amazing ability to accrue all of the journal articles and books I required.

Literature Cited Buxton, G. M., D. B. Thomas, and R. C. Froeschner. 1983. Revision of the species of the sayi-group of Chlorochloa Stål (Hemiptera: Pentatomidae). California Department of Food and Agriculture, Occasional Papers in Entomology 29: 1–25. Barton, H. E., and L. A. Lee. 1981. The Pentatomidae of Arkansas. Arkansas Academy of Science Proceedings 35: 20–25. Crevecoeur, F. F. 1905. Addition to the list of the hemipterous fauna of Kansas. Trans- actions of the Kansas Academy of Science 19: 232–237. Douce, G. K., and E. F. Suber [eds.]. 1986. Summary of insect losses from insect damage and costs of control in Georgia, 1985. University of Georgia Special Publications 55: 27. Froeschner, R. C. 1988. Family Pentatomidae Leach, 1815. The stink bugs, pp. 544–597. In T. J. Henry, and R. C. Froeschner (eds.), Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, New York. 958 pp. Hamilton, G. C. 2009. Brown marmorated stink bug. American Entomologist 55(1): 19–20. Hoffman, R. L. 2005. The Virginia species of Banasa, three decades later (Heteroptera: Pentatomidae). Banisteria 25: 41–44. Knight, H. H.1952. Review of the genus Perillus with description of a new species. (He- miptera, Pentatomidae). Annals of the Entomological Society of America 45: 229–232. Larivière, M-C. 1992. Description of Parabrochymena, new genus, and redefinition and review of Brochymena Amyot and Audinet-Serville (Hemiptera:Pentatomidae): with considerations on natural history, chorological affinities, and evolutionary relationships. Memoirs of the Entomological Society of Canada 163. pp 75. McDonald, F. J. D. 1974. Revision of the genus Holcostethus in N. America (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 82: 245–258. 218 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

McDonald, F. J. D. 1976. Revision of the genus Trichopepla (Hemiptera: Pentatomidae) in N. America. Journal of the New York Entomological Society 84: 9–22. McDonald, F. J. D. 1986. Revision of Cosmopepla Stål (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 94: 1–15. McPherson, J. E. 1982. The Pentatomoidea of Northeastern North America with Em- phasis on the Fauna of Illinois. Southern Illinois University Press, Carbondale and Edwardsville, IL, pp 240. McPherson, J. E., and R. M. McPherson. 2000. Stink Bugs of Economic Importance in America North of Mexico. CRC Press. Boca Raton, FL. pp. 253. Nielsen, A. L., and G. C. Hamilton. 2009a. Life history of the invasive species Haly- omorpha halys (Hemiptera: Pentatomidae) in northeastern United States. Annals of the Entomological Society of America 102: 608–616. Nielsen, A. L., and G. C. Hamilton. 2009b. Seasonal occurrence and impact of Haly- omorpha halys (Hemiptera: Pentatomidae) in tree fruit. Journal of Economic En- tomology 102: 1133–1140. Panizzi, A. R., J. E. McPherson, D. G. James, M. Javahery, and R. M. McPherson. 2000. Stinkbugs (Pentatomidae), pp. 421–474. In C. Schaefer, and A. R. Panizzi (eds.), Hemiptera of economic importance. Boca Raton, FL, 828 pp. Popenoe, E. A. 1885. Contributions to a knowledge of the Hemiptera-fauna of Kansas. Transactions of the Kansas Academy of Science 9: 62–64. Rider, D. A. 1986. A new species and new synonymy in the genus Tepa Rolston and McDonald (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 94: 552–558. Rider, D. A. 1995. Review of the genus Coenus Dallas with the description of C. explanatus, new species (Heteroptera: Pentatomidae). Journal of the New York Entomological Society 103: 39–47. Rider, D. A. 1996. New state records from the southeastern United States for Aelia americana (Hemiptera: Pentatomidae). Florida Entomologist 69: 251–252. Rider, D. A., and J. B. Chapin. 1992. Revision of the genus Thyanta Stål, 1862 (Het- eroptera: Pentatomidae II. North America, Central America, and the West Indies. Journal of the New York Entomological Society 100: 42–98 Rolston, L. H, 1972. The genus Menecles Stål (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 80: 234–237. Rolston, L. H. 1974. Revision of the genus Euschistus in middle America (Hemiptera, Pentatomidae, Pentatomini). Entomologica Americana 48: 1–102. Rolston, L. H., and F. J. D. McDonald. 1984. A conspectus of Pentatomini of the Western hemisphere. Part 3 (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 92: 69–86. Sailer, R. I. 1952. A review of the stinkbugs of the genus Mecidea. Proceedings of the United States National Museum 102: 471–505, plates 47–48. Schwertner, C. F., and J. Grazia. 2006. Descrição de seis espécies de Chinavia (He- miptera, Pentatomidae, Pentatominae) da América do Sul. Iheringia, Série Zoologia 96: 237–248. Thomas, D. B. 1983. Taxonomic status of the genera Chlorochloa Stål, Rhytidiloma Stål, Liodermon Kirkaldy, and Pitedia Reuter, and their included species (Hemiptera: Pentatomidae). Annals of the Entomological Society of America 76: 215–224. Thomas, D. B. 1992. Taxonomic synopsis of the asopine Pentatomidae (Heteroptera) of the western hemisphere. Entomological Society of America (Thomas Say Monograph), Lantham, Maryland. 156 pp. 2012 THE GREAT LAKES ENTOMOLOGIST 219

Thomas, D. B. 2012. Mcphersonarcys a new genus for Pentatoma aequalis Say (Heter- optera: Pentatomidae). The Great Lakes Entomologist 45: 127–133. Thomas, D. B., and T. R. Yonke. 1981. A review of the Nearctic species of the genus Banasa Stål (Hemiptera: Pentatomidae). Journal of the Kansas Entomological Society 54: 233–248. (USDA NAL) United States Department of Agriculture National Agricultural Library. 2012. Animal: species profiles: brown marmorated stink bug, May 2012. (http://www.invasivespeciesinfo.gov/animals/stinkbug.shtml) accessed 6/1/2012. Zimmer, J. T. 1910. Two new species of Pentatomidae from Nebraska. The Canadian Entomologist 42: 166–167. Zimmer, J. T. 1912. The Pentatomidae of Nebraska. University of Nebraska, Contribu- tions from the Department of Entomology, 4[1911]:1–33. 220 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 Annotated Checklist of the Pentatomidae (Heteroptera) of Connecticut Jane E. O’Donnell1 and Carl W. Schaefer1

Abstract We provide town data for the Pentatomidae in Connecticut. Although this state has been much collected, most sampling has been limited to only a few locations. Species newly recorded for Connecticut are: Halyomorpha halys (Stål), Hymenarcys nervosa (Say), Banasa euchlora Stål, B. sordida (Uhler), and Peril- lus bioculatus (Fabricius). Podisus neglectus (Westwood) may occur in the state. Other species found in neighboring states may eventually be found in Connecticut: Picromerus bidens (Linnaeus), Rhacognathus americanus Stål, Mcphersonarcys aequalis (Say), Thyanta custator custator (Fabricius), T. custator acerra McAtee, and Amaurochrous brevitylus (Barber and Sailer). We briefly analyze these data, recognizing some faunal elements. More collecting needs to be conducted in the state, so that distribution patterns outlined here can be more broadly understood, and so that species of potential conservation concern can be identified.

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Members of the Pentatomidae, or “Stink Bugs,” are ovoid robust insects of moderate to large size. The family is characterized by five-segmented antennae, presence of ocelli, a large transverse pronotum that is much broader than the head, a large prominent scutellum, the membrane of the forewing with numerous veins, tarsi three- segmented, trichobothria present on abdominal sternites four through seven but very small and inconspicuous, and the female genitalia broad and plate- like (Schuh and Slater 1995). Species are often brightly colored, and many exhibit interesting behaviors and ecological interactions. It is no wonder that Jay McPherson found a lifetime of research material studying members of this group. The junior author has known Dr. McPherson from the beginning of time, and has respected him almost from then. We are therefore pleased to offer this contribution in his honor. The limits of the family Pentatomidae and relationships within the Pen- tatomoidea are still areas of active research. Some authors treat the Scutellerinae and Acanthosomatinae not as subfamilies of the Pentatomidae but as separate families; this is the taxonomic approach that we take in this treatment, following “The Pentatomoidea Home Page” (Rider 2012). Connecticut’s fauna includes 3 of the remaining 10 pentatomid subfamilies. The North American fauna was cataloged by Froeschner (1988), who listed approximately 222 species representing 61 genera as occurring north of Mexico. Of these, 79 species in 38 genera occur in northeastern North America (United States and Canada) (McPherson 1982). McPherson (ibid.) summarized state treatments. The Connecticut fauna, treated by Parshley (1923) in Britton’s “Hemiptera of Connecticut,” included 27 genera and 45 species. The current list consists of 25 genera (not including 2 likely to occur in Connecticut but not yet reported) and 45 species (not including 4 that might eventually be found in Connecticut). The change in genus number largely reflects taxonomic changes and not distributional ones, with exceptions noted below in the checklist.

1Department of Ecology and Evolutionary Biology, Unit 3043, University of Connecti- cut, Storrs, CT 06269-3043. 2012 THE GREAT LAKES ENTOMOLOGIST 221 At the generic level, the Connecticut pentatomid fauna consists of about equal elements derived from the Neotropics, the Nearctic, and the Palearctic (Slater 1974). Within the largest subfamily, the Pentatominae, several endemic Nearctic taxa (Trichopepla, Coenus, Hymenarcys, Menecles, Dendrocoris, and Chlorochroa (Rhytidolomia)) have their closest relatives in the Neotropics (ibid.). Cosmopepla, Mormidea, Euschistus, Thyanta, Banasa and Murgantia represent the Neotropical element, with more species in each genus in the Neotropics. Slater (ibid.) interpreted Neottiglossa, Holcostethus and perhaps Chlorochroa (Chlorochroa) as Palearctic or Holarctic elements, recognizing the difficulty in distinguishing these possibilities. Brochymena and the podopine Amaurochrous are Nearctic endemics. Of the predaceous Asopinae, Rhacognathus and Picromerus, are Palearctic; Podisus, Apoecilus and Stiretrus represent Neotropical lineages, whereas Perillus is probably Nearctic (ibid.). The zoogeographic situation at the specific level is quite different. Most of the Connecticut species of Pentatomidae are strictly or almost entirely Nearctic in distribution. Only 3 species also occur in the Palearctic region and only 6 species extend substantially into the Neotropics, although our knowledge of Neotropical distributions is incomplete (ibid.).

Materials and Methods We examined over 800 specimens, some more than a century old and some collected as recently as late June 2012, from the following collections: University of Connecticut Entomology Collection (UCMS); the Connecticut Agricultural Experiment Station (CAES); Entomology Division, Peabody Museum, Yale University (YPM); and the personal collections of Chris Maier (CMC) and the junior author (CWS). The few AMNH and NMNH records cited herein were obtained from notebooks of the late James A. Slater, who gave them to the senior author many years ago. Undoubtedly more specimens have been added since he examined these collections, but it was beyond the scope of this project to obtain them. Dan Swanson (UMMZ) provided label information from specimens in the University of Michigan Museum of Zoology in emails to CWS. About half of the specimens were already authoritatively determined to species; we identified the remainder, and attached individual determination labels to each. The checklist classification at the subfamily and tribal levels follows Rider’s (2012) Pentatomoidea Home Page, except as noted, and we also used this site to verify current generic placement of species and spelling of names where possible. An * preceding the species name indicates that the species may eventually be found in Connecticut, given the proximity of known records as given in Froeschner (1988). Because we feel it will be quite a while before specimen-level data for all pentatomids are available electronically, we provide town information. Even in a small state like Connecticut, and despite the overwhelming bias of collecting localities, there is some ecological signal apparent in the distribution of stink bugs (see Discussion). Town names were standardized to the 169 names in current use; a town in brackets indicates the official town name (State of Connecticut 2012) where it differs from the name on the label. More complete label data are included for specimens representing what we believe to be the first record for the state.

Results Subfamily Asopinae Apoecilus bracteatus (Fitch); (formerly Apateticus bracteatus) Reported by Parshley (1923) from Litchfield, Prospect, Meriden, New Haven, and Madison; Parshley’s record of A. cynicus from New Canaan is referred here. Additional Connecticut records: Durham (CAES). 222 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Apoecilus cynicus (Say) Reported by Parshley (1923) from Durham, Brookfield, Portland, and Meriden; Parshley’s New Canaan record is here referred to A. bracteatus. Additional Connecticut records: South Meriden [Meriden] (CAES); Storrs [Mansfield] (UCMS); Westville [New Haven] (YPM).

Perillus bioculatus (Fabricius) Two-spotted Stink Bug. There do not appear to be any previous literature records from Connecticut. Connecticut records: Mt. Carmel [Hamden], 5 Sept. 1939, R. Beard; Mt. Carmel, Summer 1941, R. Beard; and South Meriden [Meriden], July 9, 1946, H.L. Johnson, Feeding on young Colorado potato beetle larva (all CAES).

Perillus circumcinctus Stål Reported by Parshley (1923) from Colebrook, Meriden, Milford, and Dur- ham; not reported from Connecticut in Froeschner (1988). Additional Connecticut records: New Haven (CAES).

Perillus exaptus (Say) A northern species, scarce but widespread. Reported by Parshley (1923) from Thompson, New Haven, Gales Ferry [Ledyard], Seymour, Meriden, Portland, and Wilton. Additional Connecticut records: West Willington [Willington], Mansfield, Coventry, Newington (UCMS); South Meriden [Meriden], North Stonington, Easton (YPM); Barkhamsted (CAES); Lakeville [Salisbury] (UMMZ). A specimen from East Haddam (CTM) is provisionally identified as this species. This specimen has red markings instead of the strongly contrasting white areas more typical of the species.

Perillus strigipes (Herrich-Schaeffer) (formerly Mineus strigipes) Reported by Parshley (1923) from New Canaan. Additional Connecticut records: Stratford, Shelton (CAES); Storrs [Man- sfield], Meriden, Colchester (UCMS); Woodbridge, South Meriden [Meriden], Plantsville [Southington], North Haven, North Branford, Canton, Mt. Carmel [Hamden] (YPM); Stamford (NMNH).

*Picromerus bidens (Linnaeus) Native to, and widespread in, the Palearctic; not yet reported from CT, but known from VT, MA, ME, NH and NY (Froeschner 1988), and west to Ontario in Canada (Wheeler 1999) We have seen 2 specimens from central NW Massachusetts: Heath, Frank- lin Co, 10 August 1981 (YPM). This species may eventually be found in CT.

Podisus brevispinus Phillips Note: Authorship of this species is unclear; Phillips (1982) named the species in her dissertation after discovering that the type of Arma modesta Dallas was actually a specimen of P. maculiventris; she concluded that the species referenced by McPherson and other authors as P. modestus did not actually 2012 THE GREAT LAKES ENTOMOLOGIST 223 have a name. Thomas (1992) attributes authorship to Phillips, but Rider (2012), attributes authorship to Thomas. Reported from Connecticut (as P. modestus) by Parshley (1923) from Mer- iden, Woodstock, Canaan, Kent, Lyme, Norfolk, Farmington, and New Canaan. Additional Connecticut records (some of these were as P. modestus): Corn- wall, Branford, Hamden, East Haddam (CAES); Storrs [Mansfield], Mansfield (also CWS), Cornwall, Windsor, Stonington, South Norwalk [Norwalk] (UCMS); North Stonington, Bethany, Mt. Carmel [Hamden], Union, Tunxis [Farmington], Killingworth (YPM); Putnam (AMNH); Lakeville [Salisbury] (UMMZ).

Podisus maculiventris (Say) The “Spined Soldier Bug,” is available commercially as a biocontrol agent against a wide array of agriculturally significant pests. Reported from Connecticut by Parshley (1923) from New Haven, South Glastonbury [Glastonbury], Southington, Cromwell, Windsor, Granby, Branford, Winnipauk [Norwalk], Portland, Meriden, Moosup, Pomfret, New Canaan, and Greenwich. Additional Connecticut records: Southington, Durham, Madison, Water- bury, Hamden, Stafford, Hartford, Shelton, Norfolk, North Branford (CAES); Canton, Storrs [Mansfield], Mansfield (also CWS), Coventry, Litchfield, Stafford, Gilead [Hebron], Hamden, Wallingford, Groton, West Hartford, East Haddam (UCMS); Hamden, Madison, Waterbury, Shelton, Hartford, Durham, Canton, South Meriden [Meriden], Woodbridge, Short Beach [New Haven], Stonington, No. Plain [East Haddam], North Stonington, Storrs [Mansfield], Thomaston, Guilford (YPM); Wilton, Greenwich, Litchfield (AMNH); Voluntown (CTM); Winchester (CWS); Lakeville [Salisbury] (UMMZ).

Podisus neglectus (Westwood) ( = Podisus fretus Olsen, synonmized by Thomas 1992) We are not aware of any previously published CT records. Hoffman (1971) considered the distribution “submaritime,” with records along the Atlantic coast from MA to NC; McPherson (1982) included ME, FL and the Great Lakes region of Indiana and Michigan in the range. There is a specimen from Putnam in AMNH. Putnam (Windham County), in eastern CT, which seems to fit the distribution pattern outlined above, but because we did not examine this specimen, we are hesitant to establish a new state record.

Podisus placidus Uhler Reported by Parshley (1923) from Stonington, Brookfield, New Haven, Litchfield, Wallingford, Woodbury, Portland, North Stonington, Killingly, Rain- bow [Windsor], Lyme, Ansonia, Cornwall, and Middlebury. Additional Connecticut records: Southington, Milford, Stafford, Hamden, Tolland, Waterbury, South Meriden [Meriden], Oxford, Guilford, Fairfield (CAES); Pachaug [Voluntown], Salisbury, Storrs [Mansfield], Mansfield (also CWS), Nor- wich, Meriden, Stafford, Bozrah, Willimantic [Windham] (UCMS); South Meriden [Meriden], Bethany, Savin Rock [West Haven], East Hampton, Hamden, Storrs [Mansfield], Salisbury (YPM); Stamford, New Canaan, West Cornwall [Cornwall], Putnam (AMNH); Hamden (CTM); Lakeville [Salisbury] (UMMZ).

Podisus serieventris Uhler Reported by Parshley (1923) from Branford and Woodstock. 224 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Additional Connecticut records: North Branford, Killingly, Middlebury, Orange (CAES); Willington (UCMS); vicinity Yale Forest, Windham Co. (YPM); East Had- dam, Ashford, Barkhamsted (CTM).

*Rhacognathus americanus Stål Not yet reported from CT, but known from MA, MI, IN, OH, IL, NE, MN, Alberta, and Manitoba (Froeschner 1988). It appears to be a northern species, so its presence in CT may not be all that likely.

Stiretrus anchorago (Fabricius) Reported by Parshley (1923) from Lyme. Additional Connecticut records: Shelton, Marlborough, South Meriden [Meriden], Westbrook (CAES); South Meriden [Meriden], Storrs and Mansfield Center [Mansfield], Coventry, Windsor, Stonington, Chaplin (UCMS); Mansfield (CWS); Killingworth, New Haven, Guilford, Newtown, Fairfield, Trumbull, North Haven/Wallingford line (YPM); New Canaan, Putnam (as A. fimbriatus) (AMNH); Guilford, Meriden, Salisbury (CTM).

Subfamily Pentatominae Aeliini Neottiglossa undata (Say) A northern species, ranging from southern Canada and New England west to the Pacific and south to North Carolina. Reported by Parshley (1923) from North Haven and Middlebury, and by Torre Bueno (1933) from Danbury. Additional Connecticut records: Canaan, Cromwell and New Haven (CAES); Storrs [Mansfield], Mansfield, New London, East Hartford (UCMS).

Cappaeini Halyomorpha halys (Stål) The now infamous Brown Marmorated Stink Bug, introduced from Asia to Pennsylvania in the late 1990s, made its way to Connecticut in 2008 and is now known from at least 35 towns, and in every CT county (Dr. Chris Maier, CAES, personal communication). A major agricultural pest in the mid-Atlantic states, attacking a wide variety of fruits and vegetables, as well as a nuisance insect when it enters homes to overwinter, H. halys may become the same scourge in CT as it is elsewhere.

Carpocorini Coenus delius (Say) Reported by Parshley (1923) from Scotland, New Haven, East Haven, Meriden, Thompson, Yalesville [Wallingford], Stamford, Farmington, Milford, and Marlborough. Additional Connecticut records: Gilead [Hebron], Bethany, Hamden (CAES); Storrs [Mansfield], Mansfield (also CWS), East Hartland [Hartland], George Seymour State Park [Haddam] (UCMS); Canaan, Guilford (YPM); New Canaan, Canaan, Litchfield, and Manchester (AMNH); Westport, Norwalk 2012 THE GREAT LAKES ENTOMOLOGIST 225 (NMNH). Cosmopepla bimaculata (Thomas) A common, easily recognized species, widespread in CT. Here we agree with Froeschner’s (1988) interpretation of the nomenclatural tangle involving this species name; Rider (2012) believes the correct name to be Cosmopepla lintneriana Kirkaldy. Reported by Parshley (1923) from Canaan, Thompson, Danbury, Meriden, Norfolk, Salisbury, New Haven, and Cornwall. Additional Connecticut records: Windham, Amston [Hebron], South Meriden [Meriden], Hartford, Guilford, Milford, Rainbow [Windsor], Berlin, Granby, Litchfield, Manchester, Waterbury, Stafford Springs [Stafford] (CAES); Coventry, Meriden, Plymouth, Darien, Wethersfield, “Mt. Tom State Park” [Mor- ris], Storrs and Mansfield Center [Mansfield], Mansfield (also CWS), Norwich, Stafford, Cheshire, Berlin, Bethlehem, Cromwell, Waterbury, West Hartford, Putnam, Stonington, Norfolk, Poquonock- Farmington River [Windsor], Salis- bury (UCMS); Mt. Carmel [Hamden], Washington, Plainville, North Haven, Willimantic [Windham], Hamden, Darien (YPM); New Canaan (AMNH).

Euschistus ictericus (Linnaeus) Reported by Parshley (1923) from New Haven, Branford, and Hamden. Additional Connecticut records: Durham, South Meriden [Meriden] (CAES); Storrs [Mansfield], Mansfield (also CWS), Meriden, Voluntown, Salis- bury, East Killingly [Killingly], Old Saybrook, Wequetequock [Stonington] (UCMS); Salisbury (YPM).

Euschistus politus Uhler Reported by Parshley (1923) from Portland. Additional Connecticut records: South Meriden [Meriden] (CAES).

Euschistus servus euschistoides (Vollenhoven) A more northern subspecies; large and yellowish brown, similar in color and habitus to E. variolarius. Reported by Parshley (1923) (as Euschistus euschistoides) from New Haven, Colebrook, Rockville [Vernon], Scotland, Greenwich, New Canaan, Stamford, Litchfield, Stonington, Meriden, Farmington, Darien, Derby, Milford, Northford [North Branford], Stepney [Monroe], Wilton, Cornwall, Hamden, and Marlborough; not reported from CT in Froeschner (1988). Additional Connecticut records: Portland, Wilton, Norwalk, Bethany, Wa- terbury, Prospect, North Guilford [Guilford], Monroe, Middlebury, Stonington, Northford [North Branford] (CAES); Voluntown, Norfolk, West Norfolk [Nor- folk], East Killingly [Killingly], Westford [Ashford], Cromwell, Wequetequock [Stonington], Chaplin, Phoenixville [Eastford], Beacon Falls, Union, Hamden, Coventry (UCMS); Mansfield (CWS); Portland, Middlefield, Ansonia, No. Plain [East Haddam], Woodbridge, N. Canaan [New Canaan or North Canaan], N. Haven [North Haven?], Middlefield, Portland, Canaan, Easton, Orange, Mt. Carmel [Hamden], Sharon, Southford [Southbury], South Washington [Wash- ington], Willington, North Ashford [Ashford], Woodbury (YPM); Compo Beach [Westport], Norwalk, Windsor, Stamford, South Meriden [Meriden], Groton Point [Groton], Georgetown [point where Wilton, Weston, Redding, and Ridgefield meet; or not in CT] (NMNH); Westford [Ashford], Granby, Westport (AMNH); Lakeville and Twin Lakes [Salisbury] (UMMZ). 226 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Euschistus tristigmus (Say) Reported by Parshley (1923) from Torrington, Westville [New Haven], New Canaan, East Hartford, Danbury, Manchester, Wallingford, Portland, New Haven, Colebrook, Rainbow [Windsor], Winsted [Winchester], Mystic [Stonington], Kent, Clintonville [North Branford?], and Cornwall; not reported from CT in Froeschner (1988). Additional Connecticut records: Newtown, Orange, Norwich, Sharon, Killingworth, Goshen, Salisbury, Bethany, South Meriden [Meriden] (CAES); Storrs and Mansfield Center [Mansfield], Mansfield (also CWS), Stratford, Middlefield, Tolland, Litchfield, Cobalt [East Hampton], Meriden, Somers, Stafford, Coventry, Willington, Chaplin (UCMS); East Granby, Marlborough, Granby, Hamden, Plantsville [Southington], Compo Beach [Westport], West Cornwall [Cornwall], South Meriden [Meriden], Canaan (NMNH); Guilford, Canaan, Tolland, Hamden, Meriden, North Branford (CTM); Lakeville [Salis- bury], as E. t. luridus (UMMZ).

Euschistus variolarius (Palisot de Beauvois) A large, widespread species. Reported by Parshley (1923) from Mt. Carmel [Hamden], Yalesville [Wall- ingford], New Haven, Oxford, Greenwich, Norwalk, Scotland, Hartford, New Canaan, West Haven, Westville [New Haven], Mystic [Stonington], Rainbow [Windsor], Portland, Torrington, Hamden and Wilton. Additional Connecticut Records: Salisbury, Goshen, Stafford, Woodbury, Cornwall, Branford, Windsor, Southington, Monroe, Granby, Hebron, East Hartford, Winchester, Tolland, Montville, Bethany (CAES); Portland, Salisbury, Stratford, Southington, Mansfield (also CWS), Storrs [Mansfield], Tolland, Cov- entry, Granby, Litchfield, West Willington [Willington] (UCMS); Branford, Savin Rock [West Haven], Woodbridge, Canaan, Ashford, Hartford, N. Haven [North Haven?], Killingworth (YPM); New Britain, Warehouse Point [East Windsor], Suffield, Westport, South Meriden [Meriden] (NMNH).

Holcostethus limbolarius (Stål) Reported by Parshley (1923) from New Haven, East Haven, Hamden, Manchester, Yalesville [Wallingford], Rainbow [Windsor], and Cornwall. Additional Connecticut Records: New Canaan (CAES); Storrs [Mansfield], Mansfield, Lebanon, Voluntown, Windsor, South Windsor, North Franklin [Franklin], Coventry, West Hartford, Norwich (UCMS); Seymour-Bethany- Woodbridge corner, Darien, Seymour-Woodbridge line (YPM); Meriden, South Meriden [Meriden] (NMNH).

Hymenarcys nervosa (Say) Although known from MA, NY, and RI, it has not, until now, been recorded from CT. Connecticut record: New London County, Griswold, 1 km SSW junction state route 201 and Edmund Road, 19 July 2006, leg. Chris T. Maier (CTM). *Mcphersonarcys aequalis (Say) Mcphersonarcys aequalis was placed in a new genus by Thomas (2012). Seemingly a scarce species, or at least one seldom collected; it has not yet been reported from Connecticut, but is known from MA and NY (Froeschner 1988). 2012 THE GREAT LAKES ENTOMOLOGIST 227

Menecles insertus (Say) Reported by Parshley (1923) from Wallingford, New London, and Stonington. Additional Connecticut records: Killingworth, Griswold, Kent, Hamden, South Meriden [Meriden], Branford, North Haven, New Haven (CAES); Storrs and Mansfield Center [Mansfield], Mansfield, Hamden, Danbury, Haddam, Coventry, Chaplin, Hampton (UCMS); New Haven, North Stonington, Strat- ford, Branford, North Ashford [Ashford], Willimantic [Windham], Killingworth, Bethany, No. Plain [East Haddam], Meshomasic State Forest [several towns, Middlesex and Hartford Counties] (YPM); Stamford, South Lyme [Old Lyme] (NMNH); Marlborough, on sticky band around trunk of live American white ash, Fraxinus americana L., girdled April 2005 (insect collected August 2005), Orange, East Hampton (CTM). Note: This is one of 5 species of stink bugs collected in a single UV trap on a particularly sultry late June night in eastern CT (see Banasa euchlora for location information).

Mormidea lugens (Fabricius) A common, easily recognized species. Reported by Parshley (1923) from New Haven, Mt. Carmel [Hamden], Scotland, Torrington, Branford, Danbury, West Haven, Hamden, Darien, Portland, Stonington, Milford, Kent, and North Branford, and by Torre Bueno (1933) from Danbury. Additional Connecticut Records: Sharon, Shelton, Meriden, Granby, Wil- lington, Killingworth, Marlborough, North Guilford [Guilford], East Hartford, Montville (CAES); East Killingly [Killingly], Salisbury, Phoenixville [Eastford], Voluntown, Union, West Willington [Willington], Norfolk, Wequetequock [Stonington], Westford [Ashford], Beacon Falls, West Hartford, Cromwell, Old Lyme, Rocky Hill, East Hampton, Guilford, West Norfolk [Norfolk], Plym- outh, West Ridgefield [Ridgefield], Stamford, Winsted [Winchester], Windsor, Mansfield Center [Mansfield], Mansfield (also CWS), Willimantic [Windham], Mashamoquet State Park [Pomfret], Wethersfield, West Hartford, Waterford, Hartford, Colebrook, Norwich, Willington, Bozrah, Middletown, New London (UCMS); Westville [New Haven], New Haven, Plantsville [Southington] (YPM); Killingworth, Kent, Norfolk, Columbia, North Stonington, Willington, Weston, Hamden, Seymour-Bethany-Woodbridge corner (YPM); Meriden, Canaan, Wa- terford, Orange, Southington, Burlington (CTM).

Oebalus pugnax (Fabricius) The Rice Stink Bug. Reported by Parshley (1923) from Stamford (as Solubea pugnax). Additional Connecticut Records: Madison, Norwalk (CAES); Storrs [Man- sfield] (CWS); Stratford (YPM); Hamden, on foliage of soybean (CTM).

Trichopepla semivittata (Say) Distinctively hairy, with explanate pronotal margins. Reported by Parshley (1923) from New Haven, Stony Creek [Branford], Woodmont [Milford], East Haven, and Meriden. Additional Connecticut Records: Woodbury, Bridgeport, Berlin, Stratford, Hamden, Mansfield (CAES); Beacon Falls, Storrs [Mansfield], West Hartford, Ledyard, South Meriden [Meriden], West Willington [Willington] (UCMS); North 228 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Haven, Seymour-Bethany-Woodbridge corner, Mt. Carmel [Hamden] (on Queen Ann’s Lace), Woodbridge, Long Wharf [New Haven], Sleeping Giant State Park [Hamden], Stratford, Danbury, Hamden, Orange (YPM); Westport, Greenwich, Mystic [Stonington] (NMNH); Lakeville [Salisbury] (UMMZ).

Halyini Species in this tribe are easily and often confused with the Brown Mar- morated Stink Bug.

Parabrochymena arborea (Say) Reported by Parshley (1923), as Brochymena arborea, from New Canaan, Wallingford, Lyme, New Haven, North Branford, and Stonington. Additional Connecticut Records: Westville [New Haven], Deep River, Yantic [Norwich] (CAES); Storrs and Mansfield Center [Mansfield], Mansfield, Windham Center [Windham], Windsor, Meriden, Hamden, Short Beach [Bran- ford] (UCMS); West Rock [New Haven], Middlefield, Bloomfield, North Stoning- ton, Willimantic [Windham], Fairfield, N. Haven [North Haven?], Cockaponsett State Forest [several towns, Middlesex County], No. Plain [East Haddam] (YPM); Greenwich, Putnam (AMNH); Watertown, Windsor, Cornwall, Hamden, East Hampton, Haddam, Wallingford, Hebron, Derby, Lebanon, (CTM).

Brochymena carolinensis (Westwood) Not commonly collected. Reported by Parshley (1923) from Rainbow [Windsor]. Additional Connecticut Records: Bristol (CAES); Mansfield, Storrs [Man- sfield], Danbury, New Canaan (UCMS).

Brochymena quadripustulata (Fabricius) Most common species of the genus in the state. Reported by Parshley (1923) from Westville [New Haven], New Haven, New Canaan, South Glastonbury [Glastonbury], Norwalk, Manchester, Ham- den, Middlebury, Wallingford, Westland [?doubtful CT], Bloomfield, Portland, Farmington, and Scotland. Additional Connecticut Records: Bethany, Enfield, Meriden, Milford, Wind- sor, Westport (CAES); Storrs [Mansfield], Wallingford, Stamford (UCMS); Man- sfield (CWS); Middlefield, Bridgewater, Brookfield, No. Plain [East Haddam], New Britain (YPM); Wilton, Litchfield (AMNH); Barkhamsted, Killingly (CTM) .

Nezarini Chinavia hilaris (Say) A large, nearly uniformly green species; common and widespread in CT. Reported by Parshley (1923) from Westville [New Haven], New Canaan, Poquonock [Windsor], New Haven, Lyme, Farmington, Meriden, Sharon, Middlebury, and Southington. Additional Connecticut Records: North Branford, Stamford, Norfolk, Westbrook, Portland, Stonington, West Hartford, Manchester, Harwinton, South Meriden [Meriden] (CAES); Storrs and Mansfield Center [Mansfield], Mansfield (also CWS), Stamford, Manchester, Coventry, Enfield, Vernon, West Willington [Willington], Hampton, Bethlehem, Hartford, New London, Nor- 2012 THE GREAT LAKES ENTOMOLOGIST 229 wich, Windsor (UCMS); No. Plain [East Haddam], South Meriden [Meriden], Bethany, Woodbridge, Short Beach [Branford], Manchester, Plainville, North Haven, Fair Haven [New Haven], Sleeping Giant State Park [Hamden], North Ashford [Ashford], East Rock [New Haven - Hamden line], Stonington, Willi- mantic [Windham], Seymour-Bethany-Woodbridge corner, North Stonington, Niantic [Groton], Cos Cob [Greenwich] (YPM); Stamford, Meriden, Ashford, North Branford, Guilford, Weston, Hamden, Wallingford (CTM) Note: This is another of the 5 species of stink bugs collected in the Hamp- ton, Connecticut, UV trap in June 2012.

Chinavia pennsylvanicum (Gmelin) Similar in color to C. hilaris, but smaller and more ovoid. Reported by Parshley (1923) from Stonington, who stated that there are only 3 specimens known from New England. Additional Connecticut records: We identified a single specimen from Windham County, Windham Center, 26 July 2003, J. O’Donnell, coll. (UCMS) as this species.

Chlorochroa (Chlorochroa) persimilis Horvath A scarce and local species in New England. Reported from Connecticut by Parshley (1923) from Manchester, Meriden, and Cornwall. Additional Connecticut records: Mansfield (UCMS) .

Chlorocroa (Rhytidolomia) saucia (Say) A dull olive-green to olive-brown species, inhabiting coastal marshes from MA to FL. Reported by Parshley (1923) from Greenwich, Stony Creek [Branford], New Haven, Lyme, and East Haven. Additional Connecticut Records: Madison (CAES) (UCMS) (CTM); Hamden (questionable locality, given the host plants are salt marsh grasses and this is an inland town), Savin Rock [West Haven], Groton (YPM).

Chlorochroa (Rhytidolomia) senilis (Say) Larger than C. saucia; occurs only on salt marsh grasses. Reported by Parshley (1923) from Branford, New Haven, and East Haven. Additional Connecticut records: Guilford (YPM).

Pentatomini

Banasa calva (Say) Males are easily told from B. dimiata by the shape of the pygophore opening (Hoffman 2005). Reported by Parshley (1923) from Yalesville [Wallingford]. Additional Connecticut Records: Hartford, Branford (CAES); Hampton, Middletown, Brooklyn, Windham Center [Windham], Andover, Mansfield (also CWS), Storrs [Mansfield], Willimantic [Windham], Mt. Carmel [Hamden] (UCMS); 230 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

New Haven, No. Plain [East Haddam], Willimantic [Windham], Washington, Salisbury, Bethany, Ashford (YPM); Putnam, New Canaan (AMNH); Madison, Guilford, Southington (CTM); Lakeville [Salisbury] (UMMZ). Note: This is one of 5 species of stink bugs collected in a UV trap in late June 2012 in Hampton, CT.

Banasa dimidiata (Say) The shape of the pygophore opening (males) is diagnostic (Hoffman 2005). We agree with Hoffman’s (2005) interpretation of the spelling of the name. Reported by Parshley (1923) from New Haven, Hartford, Wallingford, Portland, Stonington, Farmington, Union, Norfolk, Canaan, Southington, Hamden, and Marlborough. Additional Connecticut records: Waterbury, Middlebury, Norwalk, Corn- wall, Bethany, Monroe, Salisbury, Sharon, Goshen (CAES); Storrs and Mansfield Center [Mansfield], Mansfield (also CWS), Colebrook, Plymouth, Union, Ashford, Mt. Carmel [Hamden], Union, Ashford, North Voluntown [Voluntown], West Willington [Willington], Windsor, Windham, Hartford, Danbury (UCMS); Wil- limantic [Windham], Natchaug State Forest [Eastford] (YPM); Litchfield, New Canaan (AMNH); North Haven (CTM); Lakeville [Salisbury] (UMMZ). Note: This is one of the 5 species of stink bugs collected together in a UV trap in Hampton, CT , in late June 2012.

Banasa euchlora Stål Living on juniper, closely matching the color and pattern of the leaves. Not previously reported from CT, but listed from MA in Froeschner (1988). More data are needed before it is known whether this species was just never collected before in CT, or is truly expanding its range. CT records: Windham Co.: Hampton, Clarks Corner, 630 ft., 0.6 mi ESE BM577 power line cut, BLT [black light trap] June 20-21, 2012, E. L. Quinter, collector; an additional specimen was taken the night of July 13-14, 2012 (UCMS); Litchfield Co.: Ridgefield, 26 Chipmunk Lane, 18 January 2012, leg. Edward H. Seagraves, in wood pile (CTM). Note: This is another of the 5 species of stink bugs taken in a single UV trap in Hampton, CT, in late June 2012.

Banasa sordida (Uhler) Reported by Froeschner (1988) from Massachusetts. We are not aware of any published records for CT. Connecticut records: Tolland County: Mansfield Center [Mansfield], De- cember 21, 2011, C.W. Schaefer (1 female, UCMS); Tolland County: Mansfield Center [Mansfield], March 1, 2012, C.W. Schaefer, dead (1 male, UCMS); Wind- ham County: Willimantic [Windham], South Ridge Drive, Mercury vapor light, 9 September 2002, leg. R.J. Pupedis (1 female, UCMS); New Haven County, “N. Haven”, March 7, 1878, W.H. Patton (YPM); Middlesex County: Middletown, under powerlines between Dooley Pond and Coleman Road, 13-19 October 2005, leg. Chris T. Maier, on sticky band around girdled eastern red cedar, Juniperus virginiana L. (1 male, CTM). New Haven County: North Haven, 2 April 2001. W.L. Krinsky, on screened porch (YPM) Note: This species is easily confused with Holcostethus limbolarius. The small abdominal projection is often difficult to see inBanasa , and the overall col- 2012 THE GREAT LAKES ENTOMOLOGIST 231 oration and size of the two species are very similar. Banasa soridida tends to have greenish legs in the fresh specimens we examined; the pale lateral pronotal margins seem to be set off by darker, more thickly placed punctures in Holcostethus.

Procleticini Dendrocoris humeralis (Uhler) Reported by Parshley (1923) from Meriden, Portland, Stonington, and Killingly. Additional Connecticut records: New Haven, Bethany, South Meriden [Meriden] (CAES); Storrs [Mansfield], Hartford, Windsor, Norwich (UCMS); North Stonington, Hamden, Redding, Fairfield, Branford, New Haven, Hamden (YPM); North Haven, East Hampton (CTM).

Sciocorini Sciocoris microphthalmus Flor A small, rare, and little known species, apparently with a northern distribution. Froeschner (1988) lists it from CT. We examined no specimens.

Strachiini Murgantia histrionica (Hahn) The well-known Harlequin Bug, named for its striking, variable red and black or orange and black markings. Reported by Parshley (1923) from Meriden; he believed that this species had never become established in New England, being poorly adapted to withstand northern winters. He stated that of the two known specimens, one was brought into the state on vegetable products from the south, and the other probably so. Froeschner (1988) stated that the range of this species fluctuates markedly with the harshness or mildness of winter, migrating north during spring and summer. Froeschner (ibid.) also included New Hampshire and New York in its range. Additional Connecticut records: Mt. Carmel [Hamden], 24 August 1939, R. Beard (2 specimens); and same place and collector, 5 Sept. 1939 (all CAES).

Pentatomine genera not placed in a tribe We had difficulty identifying CTThyanta to species, despite the excellent modern treatment of Rider and Chapin (1992). They state that Thyanta calceata and Thyanta custator custator can only be reliably separated by characters of the male genitalia. We did not have enough specimens available for meaning- ful comparisons.

Thyanta calceata (Say) Reported by Parshley (1923) from New Canaan; Froeschner (1988) stated that Blatchley’s “New England” record was not confirmed by Ruckes (1957). Blatchley’s record probably came from Parshley (1923). Ruckes (1957) does not specifically exclude Connecticut, but does not include it in the list of localities for this species given in his key (p. 21); he did not deal with the species in his paper otherwise, except to note that there are no color differences between T. calceata and T. custator. Rider and Chapin (1992) listed Litchfield and New 232 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Haven counties as Connecticut localities, and their map shows several symbols in CT and neighboring Massachusetts and New York, but they provide only county records. Additional Connecticut records: Salisbury (YPM); (Twin Lakes [Salisbury] (NMNH).

*Thyanta custator accerra McAtee The range map for Thyanta custator in Ruckes (1957) includes CT; Froesch- ner (1988) listed it from CT. Thyanta accerra was treated as a separate species in Froeschner (ibid.), and reported from New York. Rider and Chapin (1992) reported Thyanta custator acerra from New York, New Jersey, New Hampshire, and Maine, so it could be found in CT. Connecticut records (provisional): several specimens from New Haven County, New Haven, Long Wharf Nature Preserve, August to October, 1994 and 1995, C.L. Remington (UCMS); New Haven County, North Haven, near exit 12 (US Hwy. 5) of Interstate 91, 41° 24.117’ N, 72° 51.039’ W, 2 Aug. 2006, leg, Chris T. Maier (CTM).

*Thyanta custator custator (Fabricius) The range map in Rider and Chapin (1992) shows a symbol in Suffolk County, NY, the closest locality to CT. This subspecies could also be in CT. Parshley (1923) thought that T. custator probably occurred in CT, because the species is common southward.

Subfamily Podopinae Podopini *Amaurochrous brevitylus (Barber and Sailer) Some old records are under the name A. parvulus. Reported in Froeschner (1988) from Massachusetts and New York, but it has not yet been taken in Connecticut.

Amaurochrous cinctipes (Say) Reported by Parshley (1923) (as Podops cinctipes) from New Haven, Me- riden, and North Branford. Additional Connecticut records: Portland, Berlin, South Meriden [Mer- iden] (CAES); Storrs and Mansfield Center [Mansfield], Mansfield (also CWS), Madison. Glastonbury, Windsor, Danbury, Stonington (UCMS); East Meriden [Meriden] (YPM).

Discussion The most striking pattern to emerge from locality information presented above is that the data are highly biased towards two centers of entomology in the state: the Connecticut Agricultural Experiment Station and Yale University in the greater New Haven area, and the University of Connecticut in Storrs. Thus, the locality data presented above are more a representation of collectors’ hometowns and favorite haunts than an accurate assessment of the ranges of species in Con- necticut. Understanding current ranges, let alone any increase, shift, or decrease in those ranges, of lesser-known insects like stink bugs will remain problematic until distribution information for many more collections is made available. It is critical to do so because conservation and land-use decisions continue to be made 2012 THE GREAT LAKES ENTOMOLOGIST 233 on a presence/absence basis of a relatively small subset of invertebrates. A few broad patterns emerge, and it is possible to recognize a number of faunal elements: 1) Coastal: this distribution pattern consists of species found along the coastal plain, often incuding the Gulf Coast. (Technically, Connecticut does not have any true coastal plain.) This is a small but well-defined element in the Connecticut fauna, and includes Thyanta custator, Chlorochroa (Rhytido- lomia) saucia, Chlorochroa (Rhytidolomia) senilis and perhaps Brochymena carolinensis. 2) Northern: This is not strictly a boreal element, but consists of species that have a distribution almost across the continent but which do not extend further south than New England, New York, northern Illinois, Nebraska, etc. This element can be considered to extend only slightly south of the old “transition” zone. Sciocoris microphthalmus would be an example. 3) North and Central: This is one of the predominant distribution patterns for Connecticut pentatomids, and includes species that are widespread in the northern states but do not extend deeply into the southern states. In the east, several species do not occur south of New Jersey, but the majority reach Virginia and North Carolina, although in some cases only in the western highlands. Some of these species are countrywide in the north, but others do not extend into or beyond the Rocky Mountains; a couple additional species are not known west of Illinois. 4) North-Central-Southern: The species that exhibit this distribution pattern are found east of the Rockies and thus have a distribution similar to 3) above, but they extend into the deep south, the majority reaching Florida and the Gulf states. 5) Country-wide: These species seem to be distributed virtually throughout the United States. 6) Southern: Only Murgantia histrionica has a distinctly southern distribution and it is actually a questionable naturally occurring species (see species account). 7) Western: Perillus bioculatus probably represents a species that originally had a strictly western distribution and then spread eastward following spread of its principal prey, the Colorado Potato Beetle. 8) Introduced (non-native): The Brown Marmorated Stink Bug, Halyomor- pha halys, was unknown in Connecticut prior to 2008.

Acknowledgments Richard Packauskas (Fort Hays State University, Kansas) and JOD started this manuscript many years ago, based on notebooks given to us by the late James A. Slater. We thank the following individuals for access and/or loan of specimens in their care: Dr. Raymond Pupedis (YPM); Dr. Chris T. Maier (CTM); and Dr. Kirby Stafford and Dr. Gale Ridge (CAES). Daniel Swanson provided the records from UMMZ. Casey James, University of Connecticut, helped transcribe and organize locality data.

Literature Cited Froeschner, R. C. 1988. Family Pentatomidae Leach, 1815. The Stink Bugs, pp. 544–597. In T. J. Henry, and R. C. Froeschner (eds.), Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden, Kobenhavn, 234 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Koln. 958 pp. Hoffman, R. L. 1971. The Insects of Virginia: No. 4: Shield Bugs (Hemiptera: Scuteller- oidea, Scutelleridae, Corimelaenidae, Cydnidae, Pentatomidae). Research Division Bulletin 67, Virginia Polytechnic Institute and State University. 61 pp. Hoffman, R. L. 2005. The Virginia species of Banasa, three decades later (Heteroptera: Pentatomidae). Banisteria 25: 41–44. McPherson, J. E. 1982. The Pentatomoidea (Hemiptera) of northeastern North America with emphasis on the fauna of Illinois. Southern Illinois University Press, Carbondale and Edwardsville. 240 pp. Parshley, H. M. 1923. Family Pentatomidae, pp. 753–776. In W. E. Britton (ed). Guide to the Insects of Connecticut. Part IV. The Hemiptera or Sucking Insects of Connecticut. Connecticut Geological and Natural History Survey of Connecticut Bulletin 34: 1–807. Phillips, K. A. 1982. A taxonomic revision of the Nearctic species of Apateticus Dallas and Podisus Herrich-Schaeffer (Heteroptera: Pentatomidae: Asopinae). Ph.D dissertation, Oregon State University. Rider, D. A. 2012. Pentatomoidea Home Page. (http://www.ndsu.nodak.edu/ndsu/rider/ Pentatomoidea/index.htm). Accessed June 2012. Rider, D. A., and J. B. Chapin. 1992. Revision of the genus Thyanta Stål, 1862 (Het- eroptera: Pentatomidae) II. North America, Central America, and the West Indies. Journal of the New York Entomological Society 100(1): 42–98. Ruckes, H. 1957. The taxonomic status and distribution of Thyanta custator (Fabricius) and Thyanta pallido-virens (Stål) (Heteroptera: Pentatomidae). American Museum Novitates 1824: 1–23. Schuh, R. T., and J. A. Slater. 1995. True Bugs of the World (Hemiptera: Heteroptera): Classification and Evolution. Cornell University Press, Ithaca, NY. Xii + 336 pp. Slater, J. A. 1974. A preliminary analysis of the derivation of the Heteroptera fauna of the Northeastern United States with special reference to the fauna of Connecticut. 25th Anniversary Memoirs, Connecticut Entomological Society, New Haven, pp. 145–213. State of Connecticut. 2012. Towns and Cities in Connecticut. (http://www.ct.gov/ ctportal/cwp/view.asp?a=843&q=257266). Accessed June 2012. Torre-Bueno, J. R. de la. 1933. Records of Connecticut Heteroptera. Bulletin of the Brooklyn Entomological Society 28(1): 36. Thomas, D. B. 1992. Taxonomic synopsis of the asopine Pentatomidae (Heteroptera) of the Western Hemisphere. Entomological Society of America, Thomas Say Foundation Monographs, Volume XVI: iv + 156 pp. Thomas, D. B. 2012. Mcphersonarcys, a new genus for Pentatoma aequalis Say (Heter- optera: Pentatomidae). The Great Lakes Entomologist 45: 127–133. Wheeler, A. G., Jr. 1999. Southern range extension of a Palearctic stink bug Picromerus bidens (Hemiptera: Pentatomidae), in North America. Entomological News 110 (2): 97–98. 2012 THE GREAT LAKES ENTOMOLOGIST 235 The Genus Sphyrocoris Mayr (Heteroptera: Scutelleridae: Pachycorinae) J. E. Eger, Jr.1,2

Abstract The genus Sphyrocoris is reviewed and found to contain a single species, S. obliquus (Germar, 1839). The following taxa are new junior synonyms of S. obliquus: Homaemus punctellus Stål, 1862; Sphyrocoris elongatus Distant, 1880; Sphyrocoris punctellus nigricans Kirkaldy, 1909; Sphyrocoris punctellus pallidomaculata Kirkaldy, 1909; and Sphyrocoris punctellus sanguineoconspersa Kirkaldy, 1909. Lectotypes are designated for H. punctellus, S. punctellus nigricans, S. punctellus pallidomaculata, and S. punctellus sanguineoconspersa.

______

The genus Sphyrocoris Mayr (1864) can be recognized among New World scutellerid genera by their small to medium size, generally cryptic brown to black mottled coloration, and an elongate ostiolar canal that is greatly expanded apically (Fig. 1). Schouteden (1904) provided a key to genera of Pachycorinae and Lattin (1964) keyed genera of scutellerids in America north of Mexico. Prior to this work, three species of Sphyrocoris were recognized: S. obliquus (Germar 1839), S. punctellus (Stål 1862) and S. elongatus Distant 1880. S. obliquus was described from Martinique and has been reported from Florida, the Caribbean, northern South America, Central America and the southwestern U.S. (Kirkaldy 1909). S. punctellus and S. elongatus were described from Mexico. Kirkaldy (1909) reported the former from Mexico, the latter from Mexico and Panama. Lattin (1964) considered the southwestern U.S. specimens to be exclusively S. punctellus and southeastern U.S. specimens to be S. obliquus, but because his work dealt exclusively with the Scutelleridae of America north of Mexico he did not treat material from Central America, South America, or the Caribbean. Lattin (1964) separated S. obliquus and S. punctellus using several characters as follows: S. obliquus - smaller, coloration relatively uniform, head narrowly rounded and convex, tylus prominent, rostrum surpassing hind coxae, scutellum narrowly rounded apically, parameres with thick, short shank; S. punctellus - larger, coloration more variable, head broadly rounded and flattened, tylus not prominent, rostrum not surpassing hind coxae, scutellum broadly rounded apically, parameres with long narrow shank. S. elongatus was not treated by Lattin because his work dealt only with taxa from America north of Mexico. Specimens of from the southwestern U.S. and the southeastern U.S. are easily separated using characters given by Lattin. However, specimens from other areas, particularly southern Central America, northern South America and the Lesser Antilles appeared to be intermediate between the two species. This study was undertaken to clarify the relationships of these species, particularly in areas south of the United States.

1Dow AgroSciences, 2608 S. Dundee Blvd., Tampa, FL 33629. 2 Research Associate, Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, FL 32602. 236 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Figure 1. Sphyrocoris obliquus metapleuron and ostiolar canal.

It is a pleasure to dedicate this paper to J. E. McPherson on the occasion of his 70th birthday in recognition of his many contributions to our knowledge of the biology and systematics of Hemiptera.

Materials and Methods Over 5,000 specimens were examined from numerous individuals and institutions. These are listed in the acknowledgments section. Specimens from various localities were evaluated based on characters used by Lattin (1964). My observations indicated that the mesosternal carinae of specimens from Mexico and the southwestern U.S. were more acutely developed and elevated posteriorly than in specimens from the southeastern U.S. (Figs. 2-3). This character was also evaluated. Approximately 175 specimens representing all geographic areas were measured and other characters evaluated. Characters evaluated and methods used are as follows: Size. overall body length and width of pronotum measurements are compared although numerous measurements were made (Table 1). Coloration. Color patterns were evaluated on a 1-5 scale as follows: 1 = Color pattern distinct and relatively sharp as in females of southeastern U.S. specimens, generally characterized by alternating bands of dark and light coloration in a vague, inverted ‘U’-shaped pattern (Fig.4). 2 = Color pattern somewhat faded, but still distinct. 2012 THE GREAT LAKES ENTOMOLOGIST 237

Figs. 2-3. Sphyrocoris obliquus. 2, prosternal carina, southwestern US; 3, Same, southeastern US. Abbreviation: C = prosternal carina. Scales = 0.5 mm.

3 = Color pattern vague, but discernible (Fig. 5). 4 = No pattern, uniformly colored. 5 = Color pattern distinct, but different from the southeastern U.S. population, characterized by 2 large black macules on the scutellum as found in some Arizona and Mexico specimens. Head shape and convexity. total head length, length before the eyes, head width immediately before the eyes, and width halfway between the eyes and the apex of the head were measured for representative specimens from the southeastern and southwestern U.S. All ratios of width vs. length were computed and analyzed to determine which best described the differences in head shape. The ratio of width immediately before the eyes to length before the eyes (anteocular ratio) showed the greatest differences between the two populations with no overlap. This ratio was then calculated for representative specimens from all geographic areas. In addition, head convexity was evaluated by measuring the width of the head immediately before the eyes and the height of the head at the same point. A ratio of width to height was calculated (convexity ratio). Length of rostrum. a ratio of rostrum length to overall body length was computed. Male genitalia. Genitalia were removed by immersing specimens in nearly boiling water for 10-15 minutes or until the pygophores of males could be 238 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Table 1. Sphyrocoris obliquus, diagnostic measurements in mm.

Males Females Mean Range Mean Range

Antennal Segment 1 0.50 0.4 - 0.6 0.54 0.4 - 0.6 Antennal Segment 2 0.51 0.4 - 0.6 0.56 0.4 - 0.7 Antennal Segment 3 0.54 0.4 - 0.7 0.59 0.4 - 0.8 Antennal Segment 4 0.80 0.6 - 1.0 0.88 0.6 - 1.2 Antennal Segment 5 0.88 0.7 - 1.1 0.94 0.8 - 1.2 Interocular Distance 1.77 1.5 - 2.2 2.01 1.7 - 2.9 Interocellar Distance 0.95 0.7 - 1.2 1.10 0.9 - 1.4 Distance from Ocelli to Eyes 0.33 0.3 - 0.4 0.37 0.3 - 0.4 Total Body Length 7.50 6.2 - 10.0 8.66 7.5 - 10.2 Head Length 1.99 1.6 - 2.4 2.17 1.8 - 2.4 Head Width 2.62 2.0 - 3.2 2.93 2.5 - 3.4 Pronotum Length 2.38 1.9 - 3.0 2.67 2.2 - 3.4 Pronotum Width 4.85 4.1 - 6.2 5.49 4.9 - 6.6 Abdomen Width 4.79 4.1 - 6.3 5.54 4.7 - 6.7 Rostral Segment 1 1.27 1.0 - 1.6 1.44 1.2 - 1.7 Rostral Segment 2 1.42 1.2 - 1.7 1.55 1.3 - 1.8 Rostral Segment 3 0.59 0.4 - 0.7 0.64 0.5 - 0.8 Rostral Segment 4 0.65 0.4 - 0.8 0.73 0.6 - 0.9 Mid-ventral Length of 2nd Abdominal Sternite 0.13 0.1 - 0.2 0.14 0.1 - 0.3 Mid-ventral Length of 3rd Abdominal Sternite 0.47 0.3 - 0.6 0.55 0.4 - 0.7 Mid-ventral Length of 4th Abdominal Sternite 0.52 0.4 - 0.7 0.61 0.5 - 0.7 Mid-ventral Length of 5th Abdominal Sternite 0.57 0.4 - 0.8 0.63 0.5 - 0.8 Mid-ventral Length of 6th Abdominal Sternite 0.56 0.4 - 0.8 0.61 0.5 - 0.8 Mid-ventral Length of 7th Abdominal Sternite 0.97 0.8 - 1.3 1.15 0.9 - 1.4 Distance from Ostiole to Legs 0.71 0.6 - 0.9 0.75 0.6 - 0.9 Distance from Ostiole to Lateral Pleural Margin 1.30 1.1 - 1.6 1.44 1.2 - 1.7 Length of Ostiolar Ruga 1.07 0.9 - 1.4 1.18 1.0 - 1.4 2012 THE GREAT LAKES ENTOMOLOGIST 239

Figures 4-5. Sphyrocoris obliquus habitus. 4) southeastern US; 5) southwestern US. Scales = 0.5 mm. 240 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 easily removed. Soft tissues were removed by immersion in hot 10% potassium hydroxide. Relative length of the shank of parameres was evaluated by measuring the length of the shank and the width of the head of the parameres. The ratios of shank length to paramere head length and body length were evaluated. Due to the availability of only a few specimens from many countries and to simplify analyses, data were grouped by larger geographical areas as follows: southwestern U.S. (California, Arizona and Texas); Mexico; Central America (Panama to the southern border of Mexico); northern South America (Colombia and Venezuela); Lesser Antilles (all Caribbean countries except those listed below under Greater Antilles); Greater Antilles (Haiti, Dominican Republic, Cuba, Puerto Rico, Jamaica and the Bahamas); southeastern U.S. (Florida, Georgia, Alabama). Measurement data and ratios across all geographic areas were analyzed using a Kruskal-Wallis test and means separated using Tukey-Kramer HSD (P = 0.05). Paired data were analyzed using a Wilcoxon test. Body coloration measurements were analyzed using ordinal logistic regression. Analytical software was JMP® Pro 9.0.3 (SAS Institute, Cary, NC). Illustrations were made using an AutoMontage® system (Syncroscopy, Cambridge, UK) through a Leica Z16 APO microscope. All measurements are given in mm. Dimensional lines in illustrations equal 0.5 mm. Label information, as it appears on the label, is presented for type specimens. When multiple labels are present, the position of the label is indicated by letters a), b), etc., with a) indicating the uppermost label.

Results and Discussion Size. The typical difference in size between females from Florida and Arizona is shown in Figs. 4-5. Relative length of males and females from different geographic areas is presented in Fig. 6. There were significant differences for both sexes (males χ2= 14.0, df = 6, P = 0.030; females χ2= 22.3, df = 6, P = 0.001). Males from the southwestern U.S. were significantly longer than those from the Greater Antilles with all other geographic areas being intermediate. Females from Mexico were significantly longer than those from Central America, the Greater Antilles, and southeastern U.S. Females from the Greater Antilles were shorter than those from the southwest U.S. and Mexico. When both sexes were combined, there were significant differences between areas χ( 2= 20.8, df = 6, P = 0.002) (Table 2). Longest specimens were from the southwestern US and the smallest from the Greater Antilles. Specimens from the southwestern U.S. were significantly longer than those from the southeastern U.S. and the Greater Antilles. Width of the pronotum showed similar trends (Table 2) with significant differences between areas χ( 2= 19.6, df = 6, P = 0.003). Specimens from the southwestern U.S. were significantly wider across the pronotum than were specimens from the southeastern U.S. and the Greater Antilles. There was a clear clinal trend toward decreased length and width as collection locality progressed from the southwestern U.S. through Mexico and Central America to South America and north through the Caribbean into the southeastern U.S. However, there was overlap in body size among most geographic areas and no significant differences between adjacent areas except that females from Mexico were longer than those from Central America. Coloration. Males and females showed differences in coloration, thus the two were analyzed separately. Analyses indicated significant differences in color rating between geographic areas (males F = 5.8, df = 6, P < 0.0001; females F = 12.8, df = 6, P < 0.0001) (Table 3). Males tended to be less strongly colored and did not show as strong a clinal trend, ranges for all geographical areas overlapping. Males from the southwest U.S. were significantly different from males collected in all areas except Mexico and Northern South America. Males from the Lesser Antilles had the lowest rating for coloration and these 2012 THE GREAT LAKES ENTOMOLOGIST 241

Figure 6. Box plots of body length for males and females of Sphyrocoris obliquus by geographic area. Abbreviations for areas are as follows: SWU = southwestern US, MEX = Mexico, CAM = Central America, SAM = South America, LAN = Lesser Antil- les, GAN = Greater Antilles, SEU = southeastern US. Box represents middle 50% of observations, vertical lines represent the upper and lower 25% of the data when these data fall outside of the box, and individual data points are outliers. Median values are indicated by the horizontal line in the box. 242 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 3.01 c 2.95 c 3.81 a 3.32 bc 3.28 bc 3.70 ab 3.56 ab Convexity Ratio = 0.05). P 1.75 c 1.93 a 1.83 b 1.64 d 1.90 a 1.82 bc 1.80 bc Anteocular Ratio 3.05 a 2.59 d 2.63 cd 2.97 ab 2.92 ab 2.83 abc 2.78 bcd Head Width 2.15 a 2.12 a 1.93 b 2.22 a 2.14 a 2.05 ab 2.04 ab Head Length 4.86 b 5.02 b 5.65 a 5.36 ab 5.30 ab 5.03 ab 5.31 ab Pronotum Width 7.47 c 8.91 a 7.81 bc 8.59 ab 8.10 abc 7.91 abc 8.25 abc Body Length Mexico Southwest US South America Lesser Antilles Greater Antilles Central America Southeastern US Geographic Area Table 2. Geographic comparisons of body length and width, head ratios width to (anteocular ratio) head width to height (head W/H ratio)*. *Means within a column with the same letter were not significantly different according to Tukey-Kramer HSD ( 2012 THE GREAT LAKES ENTOMOLOGIST 243

0 0 0 1 0 2 9 2 1 10 US 4.1 a 3.1 a Southwest = 0.05). P

0 0 2 7 1 0 0 7 4 3 3.7 a 3.9 ab Mexico Mexico .

0 0 6 0 4 7 2 0 13 13 3.7 b 2.3 b Central America Sphyrocoris obliquus

0 0 0 4 0 6 4 3 1 1 2.1 b 4.0 ab Northern S. America

1 1 4 1 0 2 7 3 0 0 2.7 c 2.1 b Lesser Antilles Geographic Area

0 0 4 0 7 3 1 0 0 11 3.7 b 1.5 b Antilles Greater

0 1 5 5 0 6 2 0 0 0 US 1.3 b 3.4 bc Southeast

Rating Table 3. Frequency of ratings for coloration in different geographic populations Males 1 2 3 4 5 Mean* Females 1 2 3 4 5 Mean* * Means in the same row with letter were not significantly different according to Nominal Logistic Regression ( 244 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 were significantly different from all other geographic areas, except southeastern U.S. Females showed a stronger clinal trend, those from the southeastern US being distinctly patterned and the pattern becoming more vague in Mexico and the southwestern U.S. Specimens from the southwestern U.S. and Mexico were significantly different from those collected in all other geographic areas, but only Mexico and southeastern U.S. specimens did not have overlap in coloration. Head shape and convexity. Head length and width, as well as anteocular ratios and convexity ratios are given in Table 2. Dorsal and lateral views of specimens from Florida and Arizona are illustrated in Figs. 7-10. There were significant differences in head length χ( 2= 25.2, df = 6, P = 0.0003) and width (χ2= 41.2, df = 6, P < 0.0001) between areas. Head length of specimens from the southeastern U.S. was significantly less than that from all other areas except the Lesser Antilles and Greater Antilles. Specimens from the southwest U.S. had significantly greater head width than specimens from the southeastern U.S. and the Lesser Antilles and Greater Antilles. The relationship of anteocular ratio to geographic locality is presented in Table 2. The mean anteocular ratio ranged from a low of 1.64 in the southeastern U.S. to a high of 1.93 in the southwestern U.S. Ranges from these two areas did not overlap. Analyses indicated significant difference between areas χ( 2= 88.7, df = 6, P < 0.0001) and there were differences in some adjacent geographic areas. Specimens from the southeastern U.S. had significantly smaller anteocular ratios than those from any other area. Anteocular ratios for specimens from the southwestern U.S. and Mexico were significantly different from those for all other areas. Despite significant differences, though, there was a clear trend toward lower ratios as you progress from the southwestern U.S. through Central America and back up to the southeastern U.S., indicating a gradual narrowing of the head. The convexity ratio also decreased across the same areas, suggesting an increase in convexity. This measure was significant across geographies χ( 2= 33.9, df = 6, P < 0.0001), with specimens from the southeastern U.S. and the Greater Antilles having significantly lower ratios than those from the southwestern U.S., Mexico and Central America. The range in anteocular ratios was greatest in the Lesser Antilles while the range in head convexity was greatest in the southwestern U.S. and the Lesser Antilles. A large range in these values for specimens from the Caribbean suggests a possible overlap of species distribution in these areas, but most values were intermediate and indicative of a single species. Length of rostrum. The ratio of rostrum length to body length showed considerable overlap among all geographic populations. The mean for this ratio ranged from 0.5 in the southwestern U.S. to 0.53 in Central and South America with no significant difference among areas (χ2= 10.3, df = 6, P = 0.113). Differ- ences in means were small and did not show a clear clinal trend. Length of the rostrum, therefore, was closely related to body size. Mesosternal carinae. The mesosternal carinae were least acutely produced and elevated posteriorly in specimens from the southeastern U.S. and the Greater Antilles. They were most acutely produced and elevated in specimens from the southwestern U.S. and Mexico. Specimens from other areas were intermediate, suggesting that there was a clinal trend similar to that seen for other characters. The greatest range was observed in specimens from northern South America. Male genitalia. Lattin (1964) stated that the male genitalia of S. obliquus and S. punctellus are very similar. He noted that the first conjunctival appendages are more slender and the sclerotized apices of the second conjunctival appendages were longer and thinner in S. punctellus. He further noted that the parameres of S. obliquus had a longer shank and more acute apex than those of S. punctellus. My examination of these characters generally supported Lattin’s observations, but differences were minor and variability in conjunctival appendages was sufficient that they could not be reliably used to separate specimens 2012 THE GREAT LAKES ENTOMOLOGIST 245

Figures 7-10. Sphyrocoris obliquus. 7-8) head, dorsal aspect; 9-10) head, lateral aspect; 7, 9) southeastern US; 8, 10) southwestern US. Scales = 0.5 mm. 246 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

from even the southeastern and southwestern U.S., let alone specimens from other areas. Parameres from representative specimens in the southwestern U.S. and southeastern U.S. were compared using the length of the shank and width of the head. For both measurements, the ranges did not overlap and means were found to be significantly different (shank χ2= 6.8, df = 1, P = 0.009; head χ2= 6.1, df = 1, P = 0.014). However, there was no difference in the ratio of paramere shaft length to paramere head width between the two geographic areas (χ2= 1.53, df = 1, P = 0.217), suggesting that differences were primarily due to size. This was further investigated by taking ratios of body length to paramere shank length and paramere head width. These ratios were not significantly different (χ2= 0.24, df = 1, P = 0.624 and χ2= 2.96, df = 1, P = 0.085, respectively). Thus, the difference in parameres between the southeastern U.S. and the southwestern U.S. appear to be primarily due to differences in body size. Although there were significant differences in some characters used by Lattin (1964) to separate S. obliquus and S. punctellus, the differences did not consistently separate populations from geographic areas outside the U.S. All characters evaluated showed a clinal trend from the southwest U.S. through Central America and northern South America, the Caribbean and into the southeastern U.S. Populations from the southeastern and southwestern US were distinct, but those from other geographic areas were consistently intermediate. Thus, there appears to be only a single species of Sphyrocoris, S. obliquus.

Taxonomic Treatment Sphyrocoris Mayr Sphyrocoris Mayr, 1864:904; Mayr, 1866:16 (keyed); Stål, 1867:495 (keyed); Schouteden, 1904:58-59 (keyed, species listed); Van Duzee, 1904:15-16; Kirkaldy, 1909:279; Van Duzee, 1917:8; Hart, 1919:168 (keyed); Blatchley, 1926:42 (keyed - Florida); Barber and Bruner, 1932:241 (keyed - Cuba); Torre-Bueno, 1939:169-170 (key to US species); Barber, 1939:278 (keyed - Puerto Rico); Lattin, 1964:161-165 (keyed, key to US species); Froeschner, 1988:691. DIAGNOSIS: Small to medium size, body convex, broadly oval, dorsum moderately punctate, fine setae usually present in punctations. Ventral punctation dense, sparse mesially on abdomen. Head triangular, broadly to narrowly rounded, slightly to moderately convex above, porrect, wider across eyes than long, moderately narrowed anterad of eyes, tylus surpassing juga by less than its width. Antennae five segmented, segments 4 and 5 slightly flattened. Scutel- lum slightly longer than wide, hemelytra exposed beyond middle of scutellum, connexiva narrowly exposed. Anterolateral pronotal margins nearly straight. Ostiolar orifices closer to metacoxae than to lateral margins of metapleura; ostiolar canal narrow basally, long, each expanded apically as large, poorly defined anteriorly directed lobe. Prosternum strongly sulcate, developed laterally into distinct carinae, mesosternum shallowly sulcate, metasternum flat or shallowly concave. Ab- dominal sterna not sulcate, lateral margins entire. Superior surfaces of tibiae sulcate. Male genital cup exposed. Proctiger simple. Aedeagus with two pairs of conjunctival appendages, first pair entirely membranous, second pair sclerotized apically. Parameres uncinate. Female genital plates simple. Dilation of spermathecal duct irregular, spermathecal pump with only proximal flange developed, spermathecal bulb digitiform. TYPE SPECIES: Pachycoris obliquus by reason of monotypy. COMMENT: The shape of the ostiolar canal (Fig. 10) is unique among New World Scutelleridae. 2012 THE GREAT LAKES ENTOMOLOGIST 247 Sphyrocoris obliquus (Germar) Pachycoris obliquus Germar, 1839:94; Herrich-Schaeffer, 1839:54-55, fig. 491; Dallas, 1851:35; Walker, 1867:48. Homaemus obliquus: Stål, 1862:81. Homaemus punctellus Stål, 1862:81-82; Walker, 1867:54. NEW SYNONYMY Sphyrocoris obliquus: Mayr, 1864:904; Stål, 1870:15; Distant, 1880:21; Schout- eden, 1904:59; Kirkaldy, 1909:279; Crawford 1913:344 (parasites); Van Duzee, 1917:8; Stoner, 1925:8-9; Blatchley, 1926:42-43 (description, host); Barber and Bruner, 1932:242-243; Torre-Bueno, 1939:169 (keyed); Barber, 1939:278; Callan, 1948:123 (host, parasites); Wolcott, 1951:190; Lattin, 1964:165-173, figs. 52-54 (keyed, described, biology); McDonald, 1966:14, 49, figs. 52-56, 429-430 (genitalia); Johnson, 1987:294-295 (egg parasite); Froeschner, 1988:691, fig. 146. Sphyrocoris punctellus: Stål, 1867:495; Stål, 1870:15; Distant, 1880:21-22, pl. 2, fig. 5; Schouteden, 1904:59, pl. 5, fig. 15; Kirkaldy, 1909:279; Van Duzee, 1917:8; Blatchley, 1934:2 (host); Torre-Bueno, 1939:169 (keyed); Bibby, 1961:329 (host); Lattin, 1964:173-180, figs. 55-57 (keyed, described, biology); Goeden and Ricker, 1976b:1171 (host); Froeschner, 1988:691; Jones, 1993:23 (hosts). Pachycoris delineatus Walker, 1867:48-49 (synonymized by Distant, 1889a:44, 51). Sphyrocoris punctellus vars. b, c, d: Stål, 1870:15. Sphyrocoris elongatus Distant, 1880:21, pl. 2, fig. 6; Distant, 1889b:314; Distant, 1893:454; Schouteden, 1904:59; Kirkaldy, 1909:279. NEW SYNONYMY. Tetyra ?delineatus: Uhler, 1886:1. Sphyrocoris punctellus nigricans Kirkaldy, 1909:279. NEW SYNONYMY. Sphyrocoris punctellus pallidomaculata Kirkaldy, 1909:279. NEW SYNONYMY. Sphyrocoris punctellus sanguineoconspersa Kirkaldy, 1909:279. NEW SYN- ONYMY. Sphaerocoris [sic!] punctellus: Van Duzee, 1923:124 (host). Sphyrocoris (Pachycoris) obliquus: Fennah, 1935:194 (host). Spyhyrocoris [sic!] obliquus: Goeden and Ricker, 1976a:925 (host). DIAGNOSIS: Coloration variable, pale to dark brown, with or without distinctive pattern; pattern typically consisting of alternating light and dark bands in an inverted ‘U’ or ‘V’ shape on the pronotum and scutellum, darker bands on the scutellum sometimes coalescing into two large macules. Coloration of head more uniform, narrow area at suture between tylus and juga darker than disc, dark coloration commonly continuing to base of head. Dorsal punctation dense, black, variably distributed, most dense on head. Venter pale, punctation concolorous to black, dense to relatively sparse. Lateral margin of abdominal sterna and most of lateral margin of propleura pale, impunctate. Legs punctate, punctures black to concolorous, sometimes black maculate. First two antennal segments pale, terminal three segments pale to dark brown. Genitalia described and figured by Lattin (1964) and McDonald (1966). TYPE MATERIAL: The location of the type of Pachycoris obliquus was reported by Lattin (1964) to be unknown and I was unable to locate it. Stål (1862) described Homaemus punctellus from Mexico without designating types. He subsequently (Stål 1870) listed three varieties (vars. b, c, and d), which 248 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Kirkaldy (1909) gave the varietal names sanguineoconspersa, pallidomaculata, and nigricans, respectively. Thus, Kirkaldy’s names were based on Stål’s specimens and because they were published before 1961, should be considered subspecies of S. punctellus. I examined five types ofHomaemus punctellus, all in the Naturhistoriska Riksmuseet, Stockholm. They were as follows: 1 male labeled: a) Mexico, b) Signt., c) Type, d) Paratypus (designated LECTOTYPE of Homaemus punctellus); 1 male labeled: a) Mexico, b) Signt., c) Paratypus; 1 female labeled: a) Mexico, b) Boucard, c) Type, d) var. b, e) Paratypus (designated LECTOTYPE of Sphyrocoris punctellus sanguineoconspersa); 1 male labeled: a) Mexico, b) Sallé, c) Type, d) var. c, e) Paratypus (designated LECTOTYPE of Sphyrocoris punctellus pallidomaculata); 1 male labeled: a) Mexico, b) Bou- card, c) Type, d) var. d, e) Paratypus (designated LECTOTYPE of Sphyrocoris punctellus nigricans). The type of Pachycoris delineatus, is a female in the British Museum, London, labeled: a) Type, b) St. Dom. 55.1, c)”17 Pachycoris delineatus”, d) Brit. Mus., Type No. Hem. 506. The type of Sphyrocoris elongatus was not located, but specimens in the British Museum from Distant’s collection, determined by Distant, leave little doubt that this is S. obliquus. DISTRIBUTION: Specimens were examined from California, Arizona, Texas, Alabama, Georgia, Florida, Cuba, Bahamas, St. Lucia, Trinidad, Bequia, Dominican Republic, Haiti, St. Vincent, Antigua, Grenada, Jamaica, Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama, Venezuela and Colombia. I have not seen specimens from Martinique, the type locality, Puerto Rico, or Barbados although it was reported from Puerto Rico by Wolcott (1951) and from Barbados by Stoner (1925). BIOLOGY: Sphyrocoris obliquus has been reported from numerous plants (Bibby 1961; Blatchley 1926, 1934; Callan 1948; Fennah 1935; Goeden and Ricker 1976a, 1976b; Jones 1993; Lattin 1964; Van Duzee 1923). I have collected all life stages from Spanish needles (Bidens bipinnata L.) in Florida and I have observed adults and nymphs feeding on the seeds of this plant. I also collected adults from Bidens sp. in Mexico. In addition, the color pattern commonly found in Florida females provides excellent camouflage on the seed heads ofB. bipinnata. This is one of the most common scutellerids in Florida and, judging by the numbers of specimens in collections, is quite common throughout much of its range. Bidens spp. are common weeds in disturbed areas and its abundance may be one reason that this insect is so common. The eggs of S. obliquus are reportedly parasitized by Trissolcus trini- dadensis Crawford (Hymenoptera: Scelionidae) (Crawford 1913, Callan 1948), a junior synonym of T. urichi Crawford (Johnson 1987).

Acknowledgments I am indebted to the following individuals and institutions for the loan and/ or donation of material to this study: J. C. Schaffner, Texas A&M University; H. Brailovsky, UNAM, Mexico; P. K. Lago, University of Mississippi; P. J. Clausen, University of Minnesota; F. Merickel, University of Idaho; W. R. Dolling and J. M. Knight, British Museum (Natural History); R. S. Zack, Washington State University; R. C. Froeschner and T. J. Henry, United States National Museum; J. D. Lattin, Oregon State University; H. D. Engleman, Salina, Kansas; L. H. Rolston, Louisiana State University; P. Dessart, J. Van Stalle, and L. Baert, Institut Royal des Sciences Naturelles de Belgique, Brussells; A. Taeger, Akademie der Landwirtschaftswissenschaften der Deutschen Demokratischen Republik, Eberswalde, Germany; J. E. Rawlins, Carnegie Museum; C. Godoy, Instituto Nacional de Biodiversidad, Costa Rica; P. Lindskog, Naturhistoriska Riksmuseet, Stockholm; P. van Doesburg, Rijksmuseum van Naturlijke Historie, Leiden; F. W. Mead, H. V. Weems and S. Halbert, Florida State Collection of Arthropods; J.-M. Maes, Museo Entomologico, Nicaragua; and C. W. and L. B. 2012 THE GREAT LAKES ENTOMOLOGIST 249 O’Brien, Florida A&M University. For the loan of material and for a review of an earlier version of this manuscript, I would like to thank D. A. Rider, North Dakota State University; D. B. Thomas, USDA, Weslaco, TX. Finally, I am grateful to P. Skelley (Florida State Collection of Arthropods) and the Florida Department of Agriculture and Consumer Services, Division of Plant Industry for assistance with and use of the AutoMontage software and associated equipment.

Literature Cited Barber, H. G. 1939. Insects of Porto Rico and the Virgin Islands - Hemiptera - Heter- optera (excluding the Miridae and Corixidae). Scientific Survey of Porto Rico 14: 263–441. Barber, H. G., and S. C. Bruner. 1932. The Cydnidae and Pentatomidae of Cuba. Jour- nal of the Department of Agriculture, University of Porto Rico 16: 231–284, pls. 24–26. Bibby, F. F. 1961. Notes on miscellaneous insects of Arizona. Journal of Economic Entomology 54: 324–333. Blatchley, W. S. 1926. Heteroptera or true bugs of Eastern North America with special reference to the faunas of Indiana and Florida. Indianapolis, IN, 1116 pp. Blatchley, W. S. 1934. Notes on a collection of Heteroptera taken in winter in the vicinity of Los Angeles, California. Transactions of the American Entomological Society 60: 1–16. Callan, E. McC. 1948. The Pentatomidae, Cydnidae and Scutelleridae of Trinidad, B. W. I. Proceedings of the Royal Entomological Society of London (B)17: 115–124. Crawford, J. C. 1913. Descriptions of new Hymenoptera, No. 8. Proceedings of the United States National Museum 46(2029): 343–352. Dallas, W. S. 1851. List of the specimens of hemipterous insects in the collection of the British Museum. London, Part 1, 368 pp. Distant, W. L. 1880. Insecta. Rhynchota. Hemiptera – Heteroptera, pp. 1–88. In F. D. Godman, and O. Salvin (eds.) Biologia Centrali - Americana. London, xx + 462 pp. Distant, W. L. 1889a. Rhynchotal notes. -Heteroptera: Scutellerinae and Graphosominae. Annals and Magazine of Natural History (7)4: 29–52. Distant, W. L. 1889b. Insecta. Rhynchota. Hemiptera – Heteroptera, pp. 305–328. In F. D. Godman, and O. Salvin (eds.) Biologia Centrali - Americana. London, xx + 462 pp. Distant, W. L. 1893. Insecta. Rhynchota. Hemiptera – Heteroptera, pp. i-xx + 329–462. In F. D. Godman and O. Salvin (eds.) Biologia Centrali - Americana. London, xx + 462 pp. Fennah, R. G. 1935. A preliminary list of the Pentatomidae of Trinidad, B. W. I. Tropi- cal Agriculture (Trinidad) 12: 192–194. Froeschner, R. C. 1988. Family Scutelleridae Leach, 1815. The shield bugs, pp. 684–693. In T. J. Henry, and R. C. Froeschner (eds.) Catalog of the Heteroptera, or true bugs, of Canada and the continental United States. New York, xix + 958 pp. Germar, E. F. 1839. Beitrage zur einer Monographie der Schildwanzen. Zeitschrift für Entomologie 1: 1–146, pl. 1. Goeden, R. D., and D. W. Ricker. 1976a. The phytophagous insect faunas of the ragweeds, Ambrosia chenopodiifolia, A. eriocentra, and A. ilicifolia, in southern California. Environmental Entomology 5: 923–930. Goeden, R. D., and D. W. Ricker. 1976b. ��The phytophagous insect fauna of the ragweed, Ambrosia psilostachya, in southern California. Environmental Entomology 5: 1169–1177. Hart, C. A. 1919. The Pentatomoidea of Illinois with keys to the Nearctic genera. Il- 250 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

linois Natural History Survey, Bulletin 13: 157–223, pls. 16–21. Herrich-Schaeffer, G. A. W. 1839. Die Wanzenartigen Insecten. Vol. 5, pp. 1–60. Johnson, N. F. 1987. Systematics of New World Trissolcus, a genus of pentatomid egg-parasites (Hymenoptera: Scelionidae): Neotropical species of the flavipes group. Journal of Natural History 21: 285–304. Jones, W. A. 1993. New host and habitat associations for some Arizona Pentatomoidea and Coreidae. Southwestern Entomologist 16(Suppl.): 1–29. Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera), vol. I, Cimicidae. Berlin, xl + 392 pp. Lattin, J. D. 1964. The Scutellerinae of America north of Mexico (Hemiptera: Heter- optera: Pentatomidae). Ph.D. Dissertation, Univ. of California, Berkeley, 350 pp. Mayr, G. L. 1864. Diagnosen neuer Hemipteren. Verhandlungen der Zoologisch- Botanischen Gesellschaft in Wien 14: 903–914. Mayr, G. L. 1866. Hemiptera in Riese der Osterreichischen Freggate Novara um die Erde in den Jahren 1857, 1858, 1859. Zoologischer Teil, II, Abt. 1. Wien, 204 pp. McDonald, F. J. D. 1966. The genitalia of North American Pentatomoidea (Hemiptera: Heteroptera). Quaestiones Entomologicae 2: 7–150. Schouteden, H. 1904. Heteroptera. Fam. Pentatomidae. Subfam. Scutellerinae. Wytsmann Genera Insectorum, fascicule 24. Brussells, 98 pp. Stål, C. 1862. Hemiptera Mexicana enumeravit speciesque novas descripsit. Stettin Entomologische Zeitung 23: 81–118. Stål, C. 1867. Bidrag till Hemipterernas systematik. Öfversigt af Kongliga Svenska Vetenskaps-Akademiens Förhandlinger 24: 491–560. Stål, C. 1870. Enumeratio Hemipterorum. 1. Enumeratio Scutellerinorum Americae. Kongeliga SvenskaVetenskaps-Akademiens Handlingar 9: 4–20. Stoner, D. 1925. Report on the Scutelleroidea collected by the Barbados-Antigua expedition from the Univ. of Iowa in 1918. University of Iowa Studies in Natural History 10(1): 1–17. Torre-Bueno, J. R. de la. 1939. A synopsis of the Hemiptera-Heteroptera of America North of Mexico. Part I, Families Scutelleridae, Cydnidae, Pentatomidae, Aradidae, Dysodiidae and Termitaphididae. Entomologica Americana 19: 141–304. Uhler, P. R. 1886. Check-list of the Hemiptera - Heteroptera of North America. Brooklyn Entomological Society, Brooklyn, i + 32 pp. Van Duzee, E. P. 1904. Annotated list of the Pentatomidae recorded from America, north of Mexico, with descriptions of some new species. Transactions of the American Entomological Society 30: 1–80. Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico,excepting the Aphididae, Coccidae and Aleurodidae. Univiversity of California Publications: Technical Bulletins. Entomology 2:i-xiv + 1–902. Van Duzee, E. P. 1923. Expedition of the California Academy of Sciences to the Gulf of California in 1921. The Hemiptera (true bugs, etc.). Proceedings of the California Academy of Sciences 12: 123–200. Walker, F. 1867. Catalogue of the specimens of Hemiptera: Heteroptera in the collection of the British Museum, London, Part. 1: 1–240. Wolcott, G. N. 1951. The Insects of Puerto Rico. Journal of the Department of ��Agricul- ture, University of Porto Rico 32[1948]: 1–975. 2012 THE GREAT LAKES ENTOMOLOGIST 251 The Stink Bugs (Hemiptera: Heteroptera: Pentatomidae) of Washington State Richard S. Zack1, Peter J. Landolt2 and Joseph E. Munyaneza2

Abstract Froeschner (1988) recorded 23 species of stink bugs (Hemiptera: Heterop- tera: Pentatomidae) from Washington State. Based on material primarily housed in the M. T. James Entomological Collection at Washington State University, the number of species is increased to 51. Three species recorded by Froeschner (1988) were not found in our collections: Apateticus crocatus (Uhler), Chloro- chroa rossiana Buxton and Thomas, and Tepa rugulosa (Say). Species recorded from Washington State for the first time are: Apoecilus bracteatus (Fitch), Perillus bioculatus (Fabricius), Podisus maculiventris (Say), P. pallens (Stål), P. placidus Uhler, P. serieventris Uhler, Zicrona caerulea (Linnaeus), Halyomorpha halys (Stål), Brochymena quadripustulata (Fabricius), B. sulcata Van Duzee, Acrosternum hilare (Say), Aelia americana Dallas, Banasa euchlora Stål, B. tumidifrons Thomas and Yonke, Chlorochroa congrua Uhler, Coenus delius (Say), Cosmopepla uhleri Montandon, Dendrocoris pini Montandon, Euschistus servus (Say), E. tristigmus (Say), E. variolarius (Palisot), Holcostethus limbolarius (Say), Neottiglossa sulcifrons Stål, N. undata (Say), Prionosoma podopioides (Uhler), Tepa yerma (Rolston), Trichopepla grossa (Van Duzee), and Amaurochrous vanduzeei Barber and Sailer.

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It is with great joy that we dedicate this paper to our friend and mentor, Jay McPherson. Jay has been an educator, researcher, colleague, and friend to all of us. He has given of his time to his science and profession but especially to the many students who, under his tutelage, have learned to be not only good scientists but also good persons. His wit and charm are unique and one cannot help but like him. His contributions through his own studies and those of his students will continue to benefit entomology, particularly those studying “bugs,” for many generations to come. In Swahili there is a title, “Mzee” which is used to describe a person who is older, advanced in years. The title however, also carries the designation of earned respect. We believe this title fits Jay very well. It is not that he has become old, but that he has become wise and lovingly respected by all who know him. Our thanks and respect, Mzee McPherson! The stink bug fauna of Washington State has been the subject of little study. Froeschner (1988) recorded 23 species of Pentatomidae from Washington, whereas numerous other species were listed from the other Pacific Northwest states of Idaho, Oregon, and Montana, as well as British Columbia, Canada. Based primarily on material in the M. T. James Entomological Collection at Washington State University, the number of species is increased to 51. Species recorded by Froeschner (1988) but not found in the collections on which this paper is based are: Apateticus crocatus (Uhler), Chlorochroa rossiana Buxton and Thomas, and Tepa rugulosa (Say). Twenty-eight species are recorded from Washington State for the first time:Apoecilus bracteatus (Fitch), Perillus

1M. T. James Entomological Collection, Washington State University, Pullman, WA 99164-6382, [email protected]. 2 Yakima Agricultural Lab., USDA, ARS, 5230 Konnowac Pass Road, Wapato, WA 98951. 252 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 bioculatus (Fabricius), Podisus maculiventris (Say), P. pallens (Stål), P. placidus Uhler, P. serieventris Uhler, Zicrona caerulea (Linnaeus), Brochymena quadripustulata (Fabricius), B. sulcata Van Duzee, Acrosternum hilare (Say), Aelia americana Dallas, Banasa euchlora Stål, B. tumidifrons Thomas and Yonke, Chlorochroa congrua Uhler, Coenus delius (Say), Cosmopepla lintneriana Kirkaldy, C. uhleri Montandon, Dendrocoris pini Montandon, Euschistus servus (Say), E. tristigmus (Say), E. variolarius (Palisot), Holcostethus limbolarius (Say), Neottiglosa sulcifrons Stål. N. undata (Say), Prionosoma podopioides (Uhler), Tepa yerma (Rolston), Trichopepla grossa (Van Duzee), and Am- aurochrous vanduzeei Barber and Sailer.

Materials and Methods This paper is based on specimens in the M. T. James Entomological Col- lection at Washington State University and the collection at the USDA, ARS Yakima Agricultural Lab located in Wapato, WA. Reported in this paper are specimens that we believe are correctly identified and were determined by credible authorities. Specimens of questionable identification are not included in the distribution records. The taxonomy of the paper is based on Froeschner (1988) for the family Pentatomidae and taxa treated in this paper are based on that work, except in cases of subsequent nomenclatural changes. In a few instances involving specimens collected in the late 19th or early 20th centuries, a generalized locality is listed on the label but a specific spot in that locality could occur in more than one county. For example, we have specimens of Apoecilus bracteatus (Fitch) from the Olympic Mountains, which span Clallam and Jefferson counties. In such cases, we have indicated the most probable county but because some uncertainty exists the county is followed by a question mark (?). Specimen records are sorted by county and site within the county; county names are capitalized. Relevant information is listed though it may not represent all information on the label(s). Dates are recorded as day, month, and year. Months are recorded by a three-letter abbreviation. In some instances, a day, month, or year are lacking on the label and thus are not shown in the records. In cases where all of the label information except one item is the same for several collecting events, we list information from the first instance and then use “same” and a change in date to represent the next record. The number of specimens represented by a single collecting event (label) is given in parenthesis after the label information. Washington State records provided by Daniel Swanson are indicated in the distributional records with the acronyms MSUC (The Albert J. Cook Arthropod Research Collection, Michigan State University Collection, East Lansing, MI) and UMMZ (University of Michigan Museum of Zoology, Ann Arbor, MI).

Results Herein we record specimens of 48 species of Pentatomidae collected from Washington State. New state records are indicated by an asterisk (*). Apateticus crocatus (Uhler), Chlorochroa rossiana Buxton and Thomas, and Tepa rugulosa (Say) were recorded by Froeschner (1988) as occurring in Washington State but are not represented in our material and are not found in the following discussion.

Subfamily Asopinae Apoecilus bracteatus (Fitch)* CLALLAM (?): Olympic Mountains (6); KING: Seattle; Pierce: Spanaway, Aug 1936, W. W. Baker, same, Sep 1936; PIERCE (?): Mt. Rainier, Aug 1895, 2012 THE GREAT LAKES ENTOMOLOGIST 253 A. L. Grant; Mt. Rainier, C. V. Piper (2); Mt. Rainier (3); STEVENS: Colville Valley, Loon Lake, 25 Jul 1882, H. M. Parshley (UMMZ); WHITMAN: Pullman, 1894, same, 12 Aug 1896 (5); same, 11 Aug 1896; same, 8-9 Aug 1896; same, 4 Oct 1929, P. Eide; same, no date; Colton, 6 Jul 1948, C. C. Shelton; YAKIMA: Yakima; 6mi SW of Tampico, 22 Oct 1911. Perillus bioculatus (Fabricius)* ADAMS: Lind, 20 Jul 1919, F. W. Carlson; BENTON: Prosser, 11 Sep 1954; CLARK: Vancouver, 25 May 1958, A. Kulin; KITTITAS: Ellensburg, no date; WHITMAN: Pullman, 16 Sep 1919, P. Putnam (UMMZ); Pullman, 27 Aug 1913; same, Apr 1917; same, 6 Jun 1918, A. L. Melander; same, Jun 1921, L. T. Qurik; same, Jun 1921, W. Johns; same, 19 May 1941, J. Beer; same, 1 Oct 1948, T. A. Thorndike; same, 9 Oct 1948, C. T. Higgins; same, 12 Oct;1948, C. F. Martinsen; same, 1948, H. Vaughn; same. 30 Aug 1949, L. R. Mewaldt; same, 18 Jul 1956; same, no date (2); same, 25 Sep 1979, R. Dingle; Wawawai, 2 May 1904; same, no date, C. V. Piper (2); YAKIMA: Moxee, 22 Apr 1909; Toppenish, 18 Jul 1923, V. Argo; same, May 1931, I. W. Bales (7). Perillus exaptus (Say) ADAMS: Sand Hills Park ca 15mi W of Washtucna, 13 Jun 1989, R. S. Zack. Podisus maculiventris (Say)* CHELAN: 7mi SSW of Wenatchee, 9 May 1981, W. J. Turner; PIERCE: Puyallup, 20 May 1931, P. Eide; WHITMAN: Pullman, Aug 2005, K. N. Landolt (6). Podisus pallens (Stål)* CHELAN: Wenatchee Heights, 26 Jun 1985. Podisus placidus Uhler* COLUMBIA: Tucannon Fish Hatchery, ca 20mi E of Dayton, 12 May 1979, W. J. Turner. Podisus servieventris Uhler* ASOTIN: Field’s Spring State Park, 4mi S of Anatone, 19 May 1973, W. J. Turner; STEVENS: Deer Lake, SE of Chewelah, 16 July1972, M. T. James; same, 1 Aug 1973; WHITMAN: Pullman, May 1907; same, May 1947. Zicrona caerulea (Linnaeus)* STEVENS: Colville, 16 Aug 1907, H. Ramstadt (UMMZ).

Subfamily Pentatominae Halyomorpha halys (Stål)* CLARK: Vancouver, 6 Oct 2010; SKAMANIA: Prindle, Riverview Road, 13 Dec 2010, K. Sheehan; YAKIMA: 1.5 mi SSE of Ahtanum Rd., 16.541°N 120.613°W, 1240 ft., 11 Aug 2012, D. Horton; same, 6 Aug 2012, P. J. Landolt, in BMSB trap. Brochymena affinis Van Duzee ASOTIN: Field’s Spring State Park, 4 mi S of Anatone, 7 Jun 1973, M. Jackson; CLARK: Jan 1968, J. T. Pennell (2); PIERCE: Parkland, 12 Feb 1952; Puyallup, 17 Sep 1940, W. W. Baker; SPOKANE: Spokane, 30 Mar 1966, R. Wiebers; same, 31 Mar 1968, J. Ulrich; same, 9 May 1968, M. Wiebers; same, 27 Jun 1968, R. Wiebers; same, 30 Aug 1968, N. Jorgensen; WHITMAN: Kamiak Butte, 24 Apr 1931, P. Eide (4); same, 26 Sep 1976, R. S. Zack; Pullman, 2 Oct 1929, P. Eide; same, 1 May 1931, R. Hansberry (4); same, Oct 1948, J. A. Hawkin; same, 31 Sep 1949, R. L. Horswill; same, 6 Apr 1977, J. Heath; Wawawai, 10 May 1959, L. Haverfield. 254 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Brochymena quadripustulata (Fabricius)* ASOTIN: Asotin, May 1923; BENTON: Hanford Site, 20 Apr 1994, R. S. Zack (2); Kennewick, Sep, 1964; Prosser, 7 Jun 1940; same, 23 May 1955, K. E. Frick; CHELAN: 1mi S of Monitor, 1 Jun 1987, R. Rathman (3); 5mi S of Wenatchee, River County Park, 11 Sep 1986, R. Rathman (10); 7mi N of Wenatchee, 8 Sep 1986, R. Rathman (5); same, 4 Jun 1987, R. Rathman; WALLA WALLA: 4mi S of Burbank 14 May 1973, W. J. Turner; WHITMAN: Almota; same, C. V. Piper; same, 27 Mar 1971, W. J. Turner; same, 7 Apr 1977, R. S. Zack; same, 19 Feb 1983, A. Mudge (2); same, 11 Mar 1985, R. S. Zack (4); Pullman, May 1932, Heinrich; same, 30 Aug 1949, L. R. Mewaldt; Wawawai; same, R. W. Doane (2); same, C. V. Piper; same, 2 Feb 1897; same, 14 May 1910; YAKIMA: Buena, 20 Oct 2011, P. J. Landolt (6); Donald, 20 May 2011, P. J. Landolt; same, 21 Oct 2011 (3); North Yakima, 28 Jul 1903, E. Jenne; Parker, 21 Oct 2011, P. J. Landolt (2); Parker Heights, 16 Mar 2001, P. J. Landolt; Toppenish, Jun 1923, V. Argo; Jul 1923; same, 4 Apr 1968, S. E. Turner; Yakima, 27 Oct 1998, P. J. Landolt; same, 2 Oct 2000; same, 3 Apr 1903; USDA Lab Near Parker, 1 Oct 1999, P. J. Landolt. Brochymena sulcata Van Duzee* Eastern Washington (unknown site);DOUGLAS: 14mi N of Wenatchee, 2 Jul 1987, R. Rathman; WHITMAN: Wawawai, 7 May 1932, I. Turner. Acrosternum hilare (Say)* ASOTIN: ca 2-3mi N of Grand Ronde River, 17 Apr 1985, R. S. Zack; CHELAN: Wenatchee, 15 May 1972, E. W. Anthon (3); Wenatchee River Site, 2 Jun 1985; 7mi N of Wenatchee, 12 Aug 1986, R. Rathman; same, 8 Sep 1986 (5); same, 4 Jun 1987; same, 31 Jul 1987; 17 Sep 1987; same, 6 Oct 1987; GAR- FIELD: 29 Sep 1942, Keene; WHITMAH: Snake River ca 1.5mi S of Wawawai, 13 Sep 1985, R. S. Zack (3); YAKIMA: Ahtanum, 4 Nov 2007, P. J. Landolt; N. Yakima, 7 Jul 1903, E. Jenne; Parker, 21 Jul 2000, P. J. Landolt; same, 21 Oct 2011 (2); Toppenish, 2 Jun 2010, P. J. Landolt; USDA Lab near Parker, 11 Aug 1999, P. J. Landolt; same, 20 Oct 1999; USDA lab near Donald, 26 Jul 2006, P. J. Landolt; Yakima, 3 Apr 1903, P. J. Landolt; Zillah, 16 May 2006, P. J. Landolt. Aelia americana Dallas* BENTON: Hanford Nuclear Site, 24 May 1995; Hanford Nuclear Site, 18-25 Apr 1998, pitfall traps, R. S. Zack and C. N. Looney; same, 1-8 May 1998 (3); same, 1-8 Jun 1998 (2); same, 8-16 June 1998; same, 18-25 May 2000. Banasa dimidiata (Say) ASOTIN: 4mi S of Anatone, 11 Jun 1982, W. J. Turner; Fields’ Spring State Park, 4mi S of Anatone, 6-7 Jun 1971, D. N. Ferro; COLUMBIA: Tucan- non Ranger Station, 20 Aug 1922, V. Argo; Tucannon Fish Hatchery, ca 13mi SE of Pomeroy, 15 May 1985, R. S. Zack; PACIFIC: Ft. Canby State Park, 11-13 Jun 1971, W. J. Turner; PIERCE: Parkland, 26 Sep 1952, V. Newhouse; SKAGIT: Rockport State Park, 7mi E of Concrete, 16-17 Jun 1971, W. J. Turner; WHITMAN: Ewan, 13 Jun 1920, R. C. Shannon (2); Pullman, 5 May 1950, J. W. Harris; same, 23 Sep 1953, O. B. Heck; same, 29 Jul 1968, R. Dawson; 22 Apr 1969; same, 1 Aug 1969; same, 3 Aug 1969; same, 27 Apr 1980, W. J. Turner; Wawawai Canyon, 15mi SW of Pullman, 15 Mar 1983, A. Mudge; Yakawawa Canyon, 7mi NW of Colton, 21 May 1977, W, J, Turner; YAKIMA: 4mi SW of Tampico, 25 Apr 2009, P. J. Landolt; 6mi SW of Tampico, 22 Oct 2011, P. J. Landolt; USDA Lab near Donald, 26 Jul 2006, P. J. Landolt; near Toppenish, 10 Aug 2010, P. J. Landolt. Banasa euchlora Stål* BENTON: Hanford Nuclear Site, 15 Sep 1994, R. S. Zack (3); YAKIMA: Parker, 24 Aug 2000, P. J. Landolt (2); USDA Lab near Donald, 20 Sep 2008, P. J. Landolt. 2012 THE GREAT LAKES ENTOMOLOGIST 255

Banasa sordida (Uhler) PIERCE: Puyallup, 22 Sep 2003, A. Antonelli; YAKIMA: Parker, 18 Oct 2011, P. J. Landolt (7); same, 19 Oct 2011 (2); USDA Lab near Parker, 13 Oct 1999, P. J. Landolt; same, 20 Oct 1999. Banasa tumidifrons Thomas and Yonke* WHITMAN: Almota, 26 Feb 1983, M. G. Means; Pullman, 6 Apr 1977, K. Raffa. Chlorochroa congrua Uhler* MASON: Lake Cushman, 13 Jul 1919, F. M. Gaige (3) (UMMZ): PACIFIC: Wallapa Bay near Nahcotta, 12 Jun 1971, W. J. Turner; SPOKANE: Cheney, 29 Apr 1969, R. Heehn. Chlorochroa granulosa (Uhler) ASOTIN: ca 2mi S of Anatone, 24 Apr 1989, R. S. Zack; 5.6mi S of Ana- tone, 24 Apr 1989, R. S. Zack; CHELAN: 7mi N of Wentachee, 8 Sep 1986, R. Rathman; DOUGLAS: 7mi S of Rock Island, 4 Sep 1986, R. Rathman; 14mi N of Wenatchee, 23 Jul 1986, R. Rathman; FERRY: 6.5mi N of Keller, 10 May 1989, R. S. Zack; LINCOLN: Davenport, 20 Jul 1987, J. Babcock (2); near Davenport, 28 Jul 2009, D. Roberts, same, 3 Aug 2009; same, 14 Jun 2010, R. Garcia; 10 Aug 2010; same, 30 Aug 2010; same, 13 Sep 2010 (2); same, 20 Sep 2010; 12mi W of Davenport, 28 Jul 1987, E. Myhre; 10.5mi S of Edwall, 9 May 1989, R. S. Zack; 3mi E of Reardan, 17 May 2010, R. Garcia; WALLA WALLA: 4mi W of Waitsburg, 16 Jul 1987, E. Myhre (2); 8mi W of Waitsburg, 16 Jul 1987, E. Myhre; WHITMAN: Steptoe Canyon 3.4mi SW of Colton, 8 May 1989, R. S. Zack; Pullman, 12 Aug 1987, R. S. Zack (2); YAKIMA: Mud Lake, 8 May 2001, C. Smithhisler; Ahtanum, 1 Oct 2011, P. J. Landolt (2): Parker, 21 Oct 2011, P. J. Landolt; near Toppenish, 1 Oct 2011, P. J. Landolt (2). Chlorochroa ligata (Say) ASOTIN: ca 2-3mi N of Grande Ronde River, 17 Apr 1985, R. S. Zack; KING: near Seattle, 10 Nov 1972 (3); LINCOLN: ca 12.2mi N of Davenport, 9 May 1989, R. S. Zack; OKANOGAN: 5.3mi E of Elmer City, 10 May 1989, R. S. Zack; PACIFIC: ca 6mi NE of Ilwaco, 22 Jun 1989, R. S. Zack; STEVENS: ca 13mi S of Fruitland, 9 May 1989, R. S. Zack (2); Chewelah, 4 Sep 1971, M. T. James (2); same, 11 Sep 1971; Deer Lake near Chewelah, 7 Oct 1972, M. T. James; WHITMAN: Pullman, 16 Sep 1982, W. J. Turner; same, 1 Feb 1983, M. G. Means; Yakawawa Canyon, 7mi NW of Colton, 21 May 1977, W. J. Turner; YAKIMA: 13 Aug 1968, B. J. Landis; same, 1 Sep 1976, F. B. Dean; same, 11 Sep 1976; same, 12 Sep 1978; same, 13 Sep 1978; Ahtanum, 6 Sep 2011, P. J. Landolt; same, 13 Sep 2008; same, 1 Oct 1911 (5); same, 7 Oct 2011 (2); same, 12 Oct 2011 (4); Near Donald, 2 Jul 2010, P. J. Landolt; 4mi SW of Tampico, 17 Aug 2011, P. J. Landolt; Parker, 21 Oct 2011, P. J. Landolt (18); Taki farm near Parker, 29 September 2011. Chlorochroa opuntiae Esselbaugh ASOTIN: Clarkston, 20 Aug 1947, C. O. Esselbaugh (2); Wilma, 16 Jul 1947, C. O. Esselbaugh; same, 20 Aug 1947. All four are paratypes. Chlorochroa sayi Stål BENTON: Prosser, E. Klostermeyer (3): same, 28 Jul 1950; same, Aug 1941, J. D. Menziescho; Richland, 16 Jul 1967, P. J. Gage; same, 20 Jul 1968; Hanford Site, 13 Jul 1967, W. Cone; same, 21 Sep 1967; same, 19 Oct 1967; CHELAN: Wenatchee, 20 May 1970, J. Brunner; Wenatchee, Squilchuck Creek, 6 Mar 1970, J. Brunner; CLARK: Vancouver, 5 Jul 1970; FRANKLIN; 3mi N of Pasco, 29 May 1957, E. F. Dailey; Moses Lake, 19 Jun 1957, E. F. Dailey; same, 17 Jul 1957, E. Stevens; GRANT: Quincy, Jones Farm, 9 Aug 1956, C. Johansen; O’Sullivan 256 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Dam, 15 Aug 1954, H. G. Davis; same, 28 Jul 1955; WHITMAN: Pullman, 6 Sep 1951, J. Ross; same, 7 Jul 1959, L. Haverfield; Smoot Hill, Pullman, 7 Aug 1970, H. S. Telford & N. Jorgensen; same, 11 Aug 1970; YAKIMA: Harrah, 24 Jul 1941, L. J. Lipovsky; Sunnyside, 28 Aug 1948, S. W. Harris; Toppenish, L. J. Lipovsky (2). Chlorochroa uhleri (Stål) ADAMS: near Hatton, 15 Aug 2008, P. J. Landolt; Lind, 25 Aug 1919, F. W. Carlson; Ritzville, 9 Sep 1920, R. C. Shannon; BENTON: Hanford Site, 30 Mar 1994, R. S. Zack (10); same, 11 Apr 1994 (3); same, 20 Apr 1994(2); same, 16 May 1994; same, 1 Jun 1994; same, 21 Jun 1994 (2): 7 Jul 1994, same, 18 Jul 1994; same, 5 Aug 1994 (2); same, 14 Sep 1994; same, 11 Oct 1994 (6); same, 25 Oct 1994 (4); COLUMBIA: near Starbuck, 22 Jul 2009; GARFIELD: sand dunes 2mi SW of Almota, 6 Jul 1989, R. S. Zack; GRANT: near Mattawa, 27 Sep 2001, P. J. Landolt; ca 11mi W of Royal City, 3 May 1989, R. S. Zack; Stratford, 17 Mar 1916; O’Sullivan Dam, 2 Jun 1954, H. G. Davis; same, 25 Sep 1954; Othello, 20 Jul 1966, D. M. Powell (4); FRANKLIN: Pasco, 4 Apr 2002, P. J. Landolt (2); LEWIS: 3 Jul 1940; LINCOLN: Davenport, 5 Jul 1987, J. Babcock (3): same, 20 Jul 1987 (2); 10mi N of Davenport, Aug 1986, J. Babcock; 12mi W of Davenport, 28 Jul 1987, E. Myhre; near Wilbur, 1 Aug 2009; same, 3 Aug 2009; PIERCE: Mt. Rainier National Park, Sunrise Ridge, 16 Aug 1921, P. Eide; SPOKANE: near Medical Lake, 8 Jul 2009; WALLA WALLA: Walla Walla, 1 May 1932, C. E. Woodworth (2); near Clyde, 5 Aug 2009; near Berryman, 9 Jul 2009 (2); 5mi S of Waitsburg, 10 Jul 1985, J. Babcock; same, 20 Jul 1986 (2); same, 27 Jul 1986 (2); same, 28 Jul 1987; WHITMAN: Almota, 29 Jul 1985, R. S. Zack (2); Pullman, 30 Jul 1894; same, 11 May 1946, C. Johansen; same, 26 Aug 1955; same, 1 Sep 1965, R. Dawson; Pullman, 13 Aug 1984; same, 12 Aug 1987, R. S. Zack; Palouse, 14 May 1963, R. Dawson; same, 1 Sep 1965; ca 15mi E of Washtucna, 3 May 1989, R. S. Zack (3); YAKIMA: Ahtanum, 28 Jun 2007, P. J. Landolt; near Ahtanum, 3 Jul 2003, P. J. Landolt; N. Yakima, 4 Aug 1903; E. Jenne; Toppenish, 28 Mar 1968, J. E. Turner; same, 8 Jul 2009, P. J. Landolt; near Toppenish, 27 Jul 2010, P. J. Landolt; ; same, 7 Aug 2010; same, 10 Aug 2010; same, 17 Aug 2010; same, 11 Aug 2011; Yakima, 17 Jul 1928, R. L. Webster; Yakima, 25 Oct 1974 (4); 5mi W of Toppenish, 15 Jun 2000, P. J. Landolt; Parker, 2 Aug 2000, P. J. Landolt; same, 24 Aug 2000 (2); same, 28 Sep 2000, same, 8 Jul 2009, same, 18 Oct 2011 (2); near Parker, 16 Aug 2001, P. J. Landolt; same, 3 Sep 2001; same, 11 Jul 2002 (2); same, 18 Jul 2002 (2); near Wapato, 17 Aug 2010, P. J. Landolt; Zillah, 29 Aug 2010, P. J. Landolt; near Zillah, 8 Jul 2009, P. J. Landolt; 15mi E of Moxee, 27 Jun 2003, P. J. Landolt; USDA Lab near Donald, 10 Aug 2004, P. J. Landolt; same, 13 Jul 2007. Codophila remota (Horvath) ADAMS: Lind, 17 Aug 1919, F. W. Carlson; 2mi N of Othello, 30 Jul 1957, E. F. Dailey (2); Washington State University Experimental Station, 18 May 2010, P. J. Landolt; BENTON: Hanford Site, 4 May 1994, R. S. Zack; same, 1 Jun 1994; same, 10 Jun 1994 (2); same, 8 Jul 1994 (2); same, 15 Jul 1994; same, 15 Sep 1994; GARFIELD: sand dunes, 2mi SW of Almota, 6 Jul 1989, R. S. Zack; LINCOLN: Davenport, 5 Jul 1989, J. Babcock; Fishtrap, 9 Aug 2011, P. J. Landolt; same, 19 Aug 2011 (2); WHITMAN: Pullman, C. V. Piper; Pullman, 8 Oct 1916; same, Oct 1931, L. Turner; Pullman, 13 Aug 1984; Farmington, 22 May 1986, E. Myhre; YAKIMA: Halvorson Farms near Toppenish, 19 May 2011, P. J. Landolt; Parker, 19 Oct 2011, P. J. Landolt; 8mi E of Moxee, 2 Jun 1957, E. F. Dailey. Coenus delius (Say)* GRANT: 5mi SE of Ephrata, R. S. Zack (3); SPOKANE: Medical Lake, 14 Jul 1920, R. C. Shannon; WHITMAN: Pullman, 29 May 1927; same, 5 Jun 1985, R. S. Zack (2). 2012 THE GREAT LAKES ENTOMOLOGIST 257

Cosmopepla bimaculata (Thomas) GRAYS HARBOR: Montessano, 13 Jun 1928, W. W. Baker; KING: Renton, Cedar River, 17 May 1952, Leuther (3); Seattle, 3 May 1939, E. E. C.; MASON: Hoodsport, 4 Aug 1969; PIERCE: Puyallup, W. H. Lawrence (3); Puyallup, 15 Sep 1927, W. W. Baker; same, 14 Oct 1927; same, 18 May 1928, same, 29 Sep 1928; same, 18 Sep 1929; Sumner, Aug 1959, E. P. Breakey; SKAGIT: 7mi E of Concrete, 16/17 Jun 1971, W. J. Turner (4); STEVENS: 31 Aug 1972 (3); WHATCOM: Bellingham, 17 Jun 1944, R. D. Shenefelt (2). Cosmopepla conspicillaris (Dallas) WHITMAN: Kamiak Butte. Cosmopepla intergressus (Uhler) CHELAN: Manson, Kinsey Ave, 0.5mi N of junction with Upper Joe Creek rd., 16 Jun 1996, collected on wild currant (50); SPOKANE: Spokane, 10 Jun 1963, N. Jorgensen; WHITMAN: Almota, 16 Apr 1983, W. J. Turner; Lyle Grove Biological area, 18 Jun 1976, W. J. Turner; 3mi W of Clarkston, 27 Apr 1985, R. S. Zack; Pullman; Pullman, 1 Jun; same, 27 May 1897; same, 23 May 1900 (4); same, 1 Jul 1901 (2); same, 10 Jul 1901; same, 28 Mar 1916; same 11 May 1932, A. Crews (4); Wawawai, 20 May 1904 (2); same, 5/21; YAKIMA: Yakima Expedition, North Yakima, 10 Jul 1903. Cosmopepla uhleri Montandon* ASOTIN: Fields’ Spring State Park, 4mi S of Anatone, 19 May 1973, D. P. Corredor; same, 19 May 1973, W. J. Turner; BENTON: Prosser, 18 May 1956, K. E. Frick; PIERCE: Electron, 25 May 1936, W. W. Baker; SPOKANE: Spokane, 9 May 1952, D. Brannon (2): WHITMAN: Pullman, 7 Jun 1948, R. R. Sanders; same, 10 Jun 1948; Wawawai, 16 May 1948. Dendrocoris pini Montandon* SPOKANE: Dartford, 1-2 Sep 1969, R. D. Gray. Euschistus conspersus Uhler ASOTIN: Fields’ Spring State Park, 4mi S of Anatone, 6-7 Jun 1971, J. A. Novak (2); CHELAN: Ardenvoir, Indian Creek, 6 Apr 1970, J. Brunner; Ardenvior, 4mi N of Mad River, 7 Jul 1973, J. Logan; Wenatchee River Site, 2 Jun 1985; Wenatchee, 15 May 1972, E. W. Anthon (4); Wenatchee, 7 June 1984; 7mi N of Wenatchee, 17 Jun 1986, R. J. Rathman; same, 12 Aug 1986 (3); same, 8 Sep 1986 (2); same, 4 Jun 1987; (5); same, 1 Jul 1987; 8mi N of Wenatchee, 18 Jun 1986, R. J. Rathman (2); same, 11 Sep 1986; 1mi S of Monitor, 18 Jun 1986, R. J. Rathman; same, 17 Jul 1986; same, 11 Aug 1987; Peshastin, Darby Canyon, G. Ching-Paulson; Columbia River Orchards, 8 Jun 1984; CLARK: Vancouver, 20 Jun 1931, P. Eide; same, 13 Oct 1935, D. Brannon; same, 17 Oct 1953, G. R. Cummings; COLUMBIA: 20mi E of Dayton, Tucannon Fish Hatch- ery, 15 May 1979, W. J. Turner; FERRY: ca 6.5mi N of Keller, 10 May 1989, R. S. Zack (2); ISLAND: Coupeville, 27 May 1898; KING: Seattle, 20 Apr 1939, E. E. C.; Renton, Cedar River, 17 May 1952, C. S. Leuthy; KITTITAS: Ellensburg; MASON: Harstine Island, 18 June 1948, G. S. Batchelor (3): PIERCE: Parkland, 20 Apr 1952, V. Newhouse, same, 6 May 1952; Fort Lewis, 4 Aug 1963, L. G. Bock; Puyallup, 10 May 1928, W. W. Baker; same, 22 Sep 2003, A. Antonelli (2); Spanaway, 14 May 1952, V. Newhouse (8); Tacoma, 12 Sep 1912; same, 9 Aug 1952, V. Newhouse; SKAGIT: 7mi E of Concrete, 16-17 Jun 1971, W. J. Turner (2): Summit Park, 11 Jun 1939, E. E. C. (2): SPOKANE: Cheney, 15 Apr 1968, R. Heehn; Dishman, 20 May 1933, R. D. Shenefelt; Spokane, 1 Jul 1968, M. Wiebers; WALLA WALLA: Walla Walla, 14 Apr 1932, C. W. Woodworth (2); same, 19 Apr 1932 (2); same, 15 May 1932; WHITMAN: Albion, 21 Feb 1973, N. Woodley (2); same, 21 Nov 1973; Almota Canyon, 16mi WSW of Pullman, 13 Jul 1989, R. S. Zack (2); Colton, 9 Jun 1948, C. C. Shelton; same, 27 Jun 258 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

1948; same 24 Jul 1948, no collector; Kamiak Butte, 14 May 1953, E. Burts; Pullman; Pullman, C. V. Piper; Pullman, Jun (2); Pullman, 9 Jul 1901; same, 1 Jun 1903; same, 2 Jun 1906; same, 2 Apr 1907; same, 25 May 1907; same, 28 May 1907; same, 29 May 1907; same, 31 May 1907; same, 2 Jun 1907; same, 3 Jun 1907; same, 17 Jun 1907; same, 5 Jun 1908; same, 15 Mar 1911; same, 19 May 1912; same, 7 May 1915; same, Jun 1921, J. A. Lowry; same, 6 May 1923, V. Argo; same, 15 May 1931, N. H. Krauss (2): same, 25 May 1931; same, 24 Apr 1932, L. Turner; same, 30 May 1940, G. Wolf; same, 22 May 1941, J. Beer; same, 25 May 1941 (2); same, 10 May 1946, G. Rogers; same, 21 May 1946, M. Hanson; same, 23 May 1948, M. T. James; same, 27 Sep 1948, N. Nullet; same, 27 Sep 1948, S. W. Harris; same, 27 Sep 1948, L. Nullet; same, 28 Sep 1948, D. N. Woodside; same, 30 Sep 1948, D. N. Woodside (2); same, 19 Aug 1949, L. R. Mewaldt; same, 17 Sep 1949, R. Dawson; same, 14 Oct 1949, G. S. Batchelor (2); same, 4 Nov 1949; same, 23 Jul 1953, A. S. Kamal; same, 6 Oct 1953, G. Trainer; same, 18 Aug 1965, R. D. Akre (2), same, 19 Aug 1965; same, 23 Aug 1965; same, 26 Aug 1965, C. Johansen (2); same, 9 Aug 1967, R. Dawson; same, 4 Sep 1968; same, 3 May 1971, J. F. McDonald; same, Aug 2005, K. Landolt; Pullman, Smoot Hill, 16 Aug 1970, H. Telford and J. Novak; same, 30 Aug 1970; Steptoe Canyon, 3.4mi SW of Colton, 8 May 1989, R. S. Zack (2); 2.5mi NE of Uniontown, 8 May 1989, R. S. Zack; Wawawai; YAKIMA: Ahtanum, 17 May 1908, P. J. Landolt, same, 1 Oct 2011, same, 12 Oct 2011 (2); near Ahtanum, 13 Jul 2007, P. J. Landolt; 2mi SW of Ahtanum, 5 Jun 2008, P. J. Landolt; Donald, 21 Oct 2011, P. J. Landolt; Parker, 29 May 2000, P. J. Landolt; same, 18 Sep 2000; same, 27 Apr 2001; same, 15 Oct 2009; near Parker, 25 May 2005, P. J. Landolt (2): Parker Heights, 4 Aug 2001, P. J. Landolt; Taki farm near Parker, 28 Jul 2006; Taki farm near Parker, 2mi N of Donald, 5 Jun 2008; same, 14 Aug 2008; USDA lab near Donald, 20 Sep 2008, P. J. Landolt; USDA lab near Parker, 6 Aug 1999, P. J. Landolt; Yakima,; Yakima, 10-2-1961 (? Month/day), B. J. Landis; N Yakima, 10 Aug 1903, E. Jenne; ca 20mi NE of Yakima, 20 Jun 1989, R. S. Zack. Euschistus servus (Say)* MASON: Lake Cushman, (UMMZ); Harstine Island, 13 Jun 1948, G. S. Batchelor; Hoodsport, 27 Aug 1966, M. T. James; PIERCE: Du Pont, 12 Jul 1947, V. Newhouse; Puyallup, 3 May 1932, J. Wilcox; SKAGIT: 7mi E of Concrete, 16-17 Jun 1971, W. J. Turner; SNOHOMISH: Everett, 25 March 1965, L. Curry; WHITMAN: Pullman, 17 Jun 1951, L. R. Cambell; WAHKIAKUM: Cathlamet, 13 Oct 1949, H. Wyrich (2). Euschistus tristigmus (Say)* CLARK: Vancouver, 14 Oct 1945, D. Brannon (4). Euschistus variolarius (Palisot)* ASOTIN: 14mi SW of Asotin, 20 Apr 1986, R. S. Zack; CHELAN: 5mi SSW of Wenatchee, 14 May 1983, W. J. Turner; CLARK: Vancouver, 13 Oct 1945, D. Brannon; LEWIS: 10mi W of Chehalis, 15 Jul 1961, C. Johansen: LINCOLN: near Davenport, 13 Sep 2010, R. Garcia; MASON: Harstine Island, 18 Jun 1948, G. S. Batchelor; STEVENS: 31 Aug 1972; WHITMAN: Almota Canyon, 16mi WSW of Pullman, 13 Jul 1989, J. back; Colton, 24 Jul 1948, C. C. Shelton; Kramer Strip near Colton, 25 June 1983, W. J. Turner (2): Steptoe Canyon, 3.4mi SW of Colton, 8 May 1989, R. S. Zack (2): Palouse, 2 Aug 1955, C. Johansen; Pullman, Apr; same, May; same, 20 May 1901; same, 8 May 1904; same, 10 May 1907, same, 12 May 1907 (3); same, 19 May 1907; same, 1 Jun 1907; same, 7 Jun 1907; same, 6 Jun 1918, A. L. Melander (3); Pullman, 29 May 1927; same, 7 May 1930; same, 17 Mar 1950, S. D. Beck; same, 23 Sep 1948, J. C. Stout; same, 30 Aug 1953, B. Okigbo; same, 25 Aug 1955, C. Johansen; same, 7 Jul 1959, L. Haverfield; same, 6 Aug 1985, J. Babcock; Pullman, Smoot Hill, 18 Jul 1970, H. Telford and J. Novak; same, 21 Aug 1970; Union Flat, 16 Jun 1916; YAKIMA: N. Yakima, 10 Jul 1903. 2012 THE GREAT LAKES ENTOMOLOGIST 259

Holcostethus abbreviatus Uhler CHELAN: Wenatchee Heights, 2 Aug 1985; GRANT: O’ Sullivan Dam, 25 Sep 1954, H. G. Davis; KLICKITAT: Bickleton, 26 Jun 1917, A. L. Melander; WHITMAN: Colton, 6 Jul 1948, C. C. Shelton (2); Pullman, no date; Pullman, 10 Jun 1948; same, 4 Jul 1894; same, 4 Jun 1907; same, 26 May 1902. Holcostethus limbolarius (Stål)* ADAMS: Ritzville, 12 Jun 1920, R. C. Shannon; Rt 26, 0.3mi E of Hopper Jct. road, 25 Sep 1989, R. S. Zack; BENTON: Rosa Unit Irrigation Exp. Station, 5 Aug 1960, E. C. Klostermeyer; Hanford, Rattlesnake Ridge, 31 Aug 1967, W. Cone; same, 19 Oct 1967; Hanford, ALE, 30 Mar 1994, R. S. Zack; Hanford, Rattlesnake Spring, 16 May 1994, R. S. Zack; CHELAN: Wenatchee, 15 May 1972, Anton (2); 1 mi S of Monitor, 14 Aug 1986, R. Rathman; same, 11 Sep 1986, R. Rathman, same, 17 Jul 1986, R. Rathman; 7 mi N of Wenatchee, 1 Jul 1987, R. Rathman; same, 12 Aug 1986; COLUMBIA: Tucannon Ranger Station, Blue Mts., 17 Aug 1922; GRANT: Gloyd Seeps, WL area, 6 mi N of Moses, 25 Sep 1989, R. S. Zack (2); LINCOLN: near Davenport, 3 Aug 2010, R. Garcia; WALLA WALLA: 5 mi S of Waitsburg, 15 Jul 1988, J. Babcock; WHITMAN: Wawawai, Jul 1900; Wawawai, 10 Oct 1949, F. E. Newby; Steptoe Cyn 9 mi W Colton, 18 Mar 1973, S. D. Berkenkamp; Steptoe Cyn 10 mi SW Colton, 22 Apr, 1982, W. S. Turner; same, 6 Apr 1985, R. S. Zack; Colton, 24 Jul 1948, C. C. Shelton; Snake River, 3 mi W of Clarkston, 27 Apr 1985, R. S. Zack; Pullman, 7 Jun 1975, P. J. Landolt; same, 11 May 1947, G. Hildreth; same, 17 Sep 1948, P. R. Ruck; same, 26 Sep 1948, R. Crossley; same, 3 May 1946, C. Johansen; same, 27 Sep 1948; same, 29 Sep 1947, F. H. Harbert; same, 23 Sep 1948, J. C. Stout; same, Oct 1912, C. Trainer; same, 24 Apr 1932, L. Turner; same, 1 May 1931, R. Hansberry; same, 13 May 1931, A. E. Bonn; same, 24 Apr 1931; same, 10 May 1931; same, 14 May 1931, P. Eide; same, 7 Jun 1978, P. J. Landolt; Spillman, Pullman, 27 Jul 1987, E. Myhre; Smoot Hill, Pullman, 27 Aug 1970, Telford & Novak; YAKIMA: Parker, 12 Jul 2000, P. J. Landolt; same, 18 Oct 2011; near Toppenish, 17 Aug 2010, P. J. Landolt; Ahtanum, 20 Sep 2010, P. J. Landolt. Holcostethus tristis (Van Duzee) WHITMAN: Albion, 21 Feb1973, N. E. Woodley (2); Colton, 6 Jul 1948, C. C. Shelton; Pullman, 21 Jan 2004, R. Crossley; same, 3 May 1904; same, 25 May 1947, L. Langdon; same, 20 Mar 1931, R. E. Bonn; Pullman, 9 Jun 1907; same, Jun 1901; same, 18 May 1904. Neottiglossa sulcifrons Stål* WHITMAN: Pullman, 4 Sep 1948, J. G. Jacobs; Pullman, 25 May 1941, J. Beer. Neottiglossa tumidifrons Downes BENTON: Hanford, ALE, 4 May 1994, R. S. Zack; Hanford, Rattlesnake Spring, 6 May 1995, R. S. Zack; Hanford Rattlesnake Ridge, 8 Jul 1994, R. S. Zack; same, 8 Jul 1994; Hanford, radio telescope, 9 Jun 1994, R. S. Zack; Han- ford, Snively Ranch, 7 Jul 1994, R. S. Zack; same, 4 May 1994 (2); WHITMAN: 4.9 mi SE of Colfax, 25 May 1989, R. S. Zack (2); Colton, 15 Jul 1960, W. Cone; Pullman, 21 May 1958; same, 9 May 1907; same, 15 May 1904; same, no date, W. H. Lawrence; Almota, 23 Apr 1983, A. Mudge; Wawawai, 12 May 1933, R. D. Shenefelt. Neottiglossa undata (Say)* COLUMBIA: 15 mi E of Dayton, Tucannon River Valley, 12 May 1973, R. A. Back; MASON: Lake Cushman, 3 Jul 1919, F. M. Gaige (UMMZ); PACIFIC: Ft. Canby SP near Ilwaco, 13-15 Jun 1973, M. Jackson; Wallapa Bay near Nahcotta, 12 Jun 1971, W. J. Turner (3); PIERCE: Parkland, 21 Apr1952, V. Newhouse 260 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

(3); Ft. Lewis, 4 Aug 1963, L. G. Bock; WHITMAN: Almota, 26 Apr 1931, A. L. Grant; Pullman, 22 May 1941, J. Beer; same, 5 Jun1985, R. S. Zack; same, May 1947, same, 25 May 1902; same, 30 May1904; same 9 May 1907, (2); same, 21 Apr 1907; same, 21 May 1907; same, 21 May 1907; same, 17 May 1916; same, 7 May 1908; same, 4 May 1929; south of Pullman, 18 Jun 1950, R. B. Spurrier. Prionosoma podopioides (Uhler)* CHELAN: Wenatchee, Apr 1912; Chelan, 27 Apr 1912, M. A. Yothers; DOUGLAS: Moses Coulee, 6 May 1933; GARFIELD: Snake River, sand dunes, ca 2 mi SW of Almota , 6 Jul 1989, R. S. Zack (2); GRANT: Grand Coulee, Steamboat Rock, 10 Jun 1902; KITTITAS: Vantage, 10 May 1936, W. W. Baker. Tepa yerma (Rolston)* ADAMS: Othello, 15 Aug 1949, E. Klostermeyer (MSUC); CHELAN: Wenatchee, 9 Sep 1940, (2). Thyanta pallidovirens (Stål) ASOTIN: 2-3 mi N of Grand Ronde River, 14 Apr 1985, R. S. Zack; 2 mi S of Asotin, 24 Apr 1989, R. S. Zack; Anatone, 2 Oct 1949, L. R. Mewaldt; 4 mi S of Anatone, 30 Apr 1977, K. Raffa; 7.2 mi S of Anatone, 24 Apr 1989, R. S. Zack; 10 mi S of Anatone, 24 Apr 1989, R. S. Zack; BENTON: Richland, 12 Jun 1969, P. J. Gage; same, 12 Jul 1969; Prosser, Rosa Unit, IAREC, 26 Aug 1958; Hanford, ALE, 30 Mar 1994, R. S. Zack; same, 11 Apr 1994 (6); same, 20 Apr 1994 (4); same, 16 May 1994; same, 25 Oct 1994 (2); Hanford, Benson Spring, 30 Mar 1994, R. S. Zack; Hanford, radio telescope, 9 Jun 1994, R. S. Zack (16); same, 18 Jul 1994; Hanford, Rattlesnake Ridge, 8 Jul 1994, R. S. Zack (5); same, 18 Jul 1994 (3); same, 12 Aug 1994 (6); same, 15 Sep 1994; Hanford, Rattlesnake Spring, 3 May 1994, R. S. Zack; same, 21 Jun 1994 (2); same, 11 Oct 1994; Hanford, Lower Snively Spring, 1 Jun 1994, R. S. Zack (3); Hanford, Snively Ranch, 4 May 1994, R. S. Zack (2); Hanford Snively Spring, 4 May 1994, R. S. Zack; CHELAN: Wenatchee Heights, 2 Aug 1985, R. Rathman (2); Wenatchee, Jun 1940; same, 24 Jun 1940 (2); same, 28 Jun 1940; 7 mi N of Wenatchee, 1 Jul 1987, R. Rathman (6); same, 14 Jul 1986 (2); same, 12 Aug 1986; same, 8 Sep 1986 (4); 1 mi S of Monitor, 14 Aug 1986, R. Rathman; same, 11 Sep 1986; same, 28 Jul 1987; same, 16 Sep 1987; Cashmere, 30 Apr 1939, E. C. C. (2); CLARK: Vancouver, 13 Aug 1970; FERRY: 7.7 mi S of Keller, 10 May 1989, R. S. Zack (5); GARFIELD: 2 mi SW of Almota, 6 Jul 1989, R. S. Zack (8); 14 mi E of Pomeroy, 22 Apr 1988, R. S. Zack (2); 11 mi SSW of Pomeroy, 26 Apr 1988, R. S. Zack; GRANT: Gloyd Seeps WL area, 6 mi N of Moses Lake, 25 Sep 1989, R. S. Zack (3); Mattawa, 17 Sept 2001, Adams and Green; same, 27 Sep 2001, P. J. Landolt; near Mattawa, 17 Sep 2001, P. J. Landolt; KING: Seattle, 17 Jun 1930; KITTITAS: 5 mi S of Ellensburg, 13 Jun 1989, R. S. Zack; 6 mi W of Vantage, 3 May 1989, R. S. Zack (5); KLICKITAT: 8.5 mi ENE of Wishram, 15 Apr 1988, R. S. Zack; LINCOLN: 10.5 mi S of Edwall, 9 May 1989, R. S. Zack (2); 12.2 mi N of Davenport, 9 May 1989, R. S. Zack (2); OKANOGAN: Malott, Jul 1938, E. C. C.; 2.1 mi E of Elmer City, 10 May 1989, R. S. Zack (3); 1.5 mi N of Elmer City, 10 May 1989, R. S. Zack; 5.3 mi E of Elmer City, 10 May 1989, R. S. Zack (4); 8.4 mi E of Elmer City, 10 May 1989, R. S. Zack; SPOKANE: Spokane, 27 Jul 1968, M. Wiebers; STEVENS: 13 mi S of Fruitland, 9 May 1989, R. S. Zack; WALLA WALLA:5 mi S of Waitsburg, 15 Jul 1988, J. Babcock (3); WHITMAN: 15 mi E of Washtucna, 3 May 1989, R. S. Zack; 13.5 mi N of Wilbur, 10 May 1989, R. S. Zack (2); Steptoe Cyn, 3.4 mi SW of Colton, 8 May 1989, R. S. Zack; Steptoe Cyn 10 mi SW of Colton, 29 Jul 1976, W. J. Turner; same,16 Sep 1982 (4); Almota, 10 Sep 1975, W. S. Turner; same, 19 Feb 1983, A. Mudge (3); same, 13 Apr 1985, W. S. Turner; Pullman, 12 Aug 1984; same, 13 Aug 1984, (2); same, 8-9 Aug 1953, B. Okigbo; same, 8 Oct 1949, G. S. Batchelor; same, 11 Nov 1949 (2); same, Nov 1949, G. S. Batchelor; same, 9 Aug 2012 THE GREAT LAKES ENTOMOLOGIST 261 1967, B. A. Freeman; same, 13 Oct 1947, D. D. Earp; Kamiak Butte, Pullman, 14 May 1953, E. C. Trivette; Lyle Grove Biological area, 8 mi SW of Pullman, 13 Aug 1983, R. S. Zack; Wawawai, 10 Oct 1949, F. E. Newby, same, 13 Oct 1948, H. L. Latimer (2); same, 13 Oct 1947, C. C. Rosecrans; same, 3 Oct 1948, J.C. Stout (3); same, 2 Oct 1953, O. B. Heck (3); YAKIMA: 28 mi S of Toppenish, 25 Apr 1989, R. S. Zack (2); Yakima, 30 Aug 1930, A. R. Rolfs; same, 25 Oct 1974, (2); same, 14 Jul 2008, P. J. Landolt; 6 mi SW Tampico, 22 Oct 2011, P. J. Landolt; USDA lab near Parker, 7 Jul 1999, P. J. Landolt,; same, 1 Jul 1999, same ,4 Aug 1999 (2); Parker, 21 Jul 2000, P. J. Landolt (2); same, 18 Sep 2000; same, 21 Oct 2011 (4); same, 19 Oct 2011 (5); same, 18 Oct 2011 (3); near Parker, 16 Aug 2001, P. J. Landolt; Parker Heights, 21 Aug 1999, P. J. Landolt; 15 mi E of Moxee, 19 Jul 2000, P. J. Landolt; same, 25 Jun 2003 (2); same, 27 Jun 2003 (2); Terrace Heights, 21 Jul 2011, P. J. Landolt; near Donald – USDA lab, 26 Jul 2006, P. J. Landolt (5); same, 5 Aug 2009; same, 10 Aug 2004; near Donald, 27 Aug 2007, P. J. Landolt; near Toppenish, 27 Jul 2010, P. J. Landolt; same, 10 Aug 2010; Buena, 20 Oct 2011, P. J. Landolt. Trichopepla aurora Van Duzee WHITMAN: Pullman, 6 Jun 1918, A. L. Melander. Trichopepla dubia (Dallas) COLUMBIA: Tucannon Ranger Station, Blue Mts, 20 Aug 1922, N. Argo; PIERCE: Mt. Rainer, Paradise Park, Aug 1917, A. L. Melander (2); WHITMAN: Pullman, 10 Jun 1907. Trichopepla grossa (Van Duzee)* BENTON: Hanford, radio telescope, 9 Jun 1994, R. S. Zack; WHITMAN: Pullman, 25 May 1902; same, 27 Aug, 1913; same, 7 May 1930; same, 15 May 1931, R. Hansberry; same, 24 Apr 1932, L. Turner (2).

Subfamily Podopinae Amaurochrous vanduzeei Barber and Sailer* BENTON: Hanford Nuclear Site; West Lake, N46°36.066’ W119°32.788’, 4-18 Apr 1998, C. Looney, pitfall traps (3); same, 23 May – 1 Jun 1998 (10); same, 1-8 Jun 1998 (3); same, 8-16 Jun 1998 (3); same, 9-24 Oct; same, 25 Nov – 17 Dec 1998; same, 5-12 Mar 1999 (3), same, 19-26 Mar 1999 (4); same, 26 Mar – 2 Apr 1999 (3); same, 2-9 Apr 1999 (2); same, 16-24 Apr 1999 (3); same, 24-30 Apr 1999; same, 18-25 Jun 1999; CHELAN: 1mi S of Monitor, 17 Jul 1986; GARFIELD: sand dunes, 4mi NW of Lower Granite Dam, 25 Sep 1985, R. S. Zack (4); GRANT: Soda Lake near O’Sullivan Dam, 2 May 1982, W. J. Turner (2); Lake Lenore, 8 Jul 1993, R. S. Zack; YAKIMA: 8mi SW of Tieton, 11-12 Jun 1973, M. Jackson.

Discussion We increase the number of species of Pentatomidae recorded from Wash- ington State from 23 (Froeschner 1988) to 51. Washington is an extremely diverse state in terms of environment from coastal rain forest on the Olympic Peninsula to dry desert in central Washington to alpine and extensive forests. While the state has been subject to extensive urbanization and agriculture, large tracts of native and undisturbed habitat remain. Our collection records represent material collected primarily in eastern Washington (east of the Cascade Mountains) as Washington State University is located at the extreme eastern edge of the state where most collecting by university and collection personnel has taken place. Specific collecting for stink bugs in other areas of Washington will probably show that many species are more widely distributed than our records indicate and that other species occur in the state. 262 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Acknowledgments We want to thank Daniel Swanson at the University of Michigan Museum of Zoology for providing specimen records from the University of Michigan Museum of Zoology and The Albert J. Cook Arthropod Research Collection at Michigan State University. Michael R. Bush, Washington State University Extension, provided a helpful review of the manuscript. Chris Looney, Wash- ington State Department of Agriculture, and Todd Murray, Washington State University Extension, provided records of the recently introduced species, Halyomorpha halys.

Literature Cited Froeschner, R. C. 1988. Family Pentatomidae Leach, 1815, pp. 544–597. In: T. J. Henry, and R. C. Froeschner (eds.), Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, New York. 2012 THE GREAT LAKES ENTOMOLOGIST 263 An updated synopsis of the Pentatomoidea (Heteroptera) of Michigan. D. R. Swanson1

Abstract An overview of the 75 species of Pentatomoidea found in Michigan is presented, along with updated identification keys, distributional lists, and relevant literature. New state records for Corimelaena agrella McAtee (Thyreocoridae), Melanaethus robustus Uhler (Cydnidae), Pangaeus bilineatus (Say) (Cydnidae), and Banasa sordida (Uhler) (Pentatomidae: Pentatominae) are included.

______

The knowledge of the Pentatomoidea in Michigan is probably the greatest for any group of terrestrial Heteroptera in the state. O’Brien (1983, 1988) listed the relevant sources of information for the terrestrial arthropods of Michigan. Townsend (1980) and Hussey (1921, 1922a) contributed to the beginnings of the knowledge of the Pentatomoidea of Michigan, cataloging the Heteroptera found in the vicinity of Constantine, Saint Joseph County and Berrien County, respectively. Stoner (1922) also treated the species encountered around a biological station in the northernmost Lower Peninsula (Cheboygan County). McPherson (1970, 1979b) greatly augmented the knowledge and the known diversity of the Michigan fauna. In addition to these state-focused studies, the majority of Michigan’s taxa are known from the surrounding area, and many extra-Michiganian references have supplied applicable information. As a tribute to Dr. J. E. McPherson on his 70th birthday and retirement, for his great and numerous contributions to the field of pentatomology, I am pleased to present herein an update of his landmark study of the Pentatomoidea of Michigan.

Materials and Methods The author examined the pentatomoid holdings of the two major university collections in southern Michigan. County records were compiled, identification keys were modified, and the existing natural history and systematic information, both Michiganian and some extralimital, was summarized. The identification of the 8450 specimens included in this study was rendered or confirmed by the author, although the vast majority of specimens already had been studied by McPherson (1970, 1979b). All specimens reside in one of the collections listed below unless otherwise noted. Collection dates indicate the earliest and latest adults examined and refer specifically to specimens collected in Michigan. In the few instances where it is provided, label data are not transcribed verbatim but the locality information is included in its entirety. I have eschewed the use of individual species distribution maps in this synopsis owing to the lack of significant change from the maps provided by McPherson (1970) as well as the large number of species included in this treatment; therefore, the distributions are provided at the end of each species account as a list of counties. Locations of Michigan counties from which specimens were collected are depicted in Figure 1.

13111 Scenic Lake Dr. Apt. 34, Ann Arbor, MI 48108. (email: [email protected]). 264 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Figure 1. The counties of the State of Michigan.

As this study is an update of the extensive work done by McPherson (1970, 1979b), I have endeavored mostly to expand and supplement that work. Since that time, one species (Halyomorpha halys (Stål)), in addition to the four reported here, have been added to the Michigan fauna. Two hundred seventy four county records have been added. Additionally, much of the taxonomy has changed in the past forty years; thus, references to more contemporary systematic treatments are provided, and 14 nomenclatural changes affecting 15 species and 1 family-group have been incorporated (noted within the relevant section). The number of additions and changes necessitated that new dichotomous keys to the Michigan fauna be synthesized. 2012 THE GREAT LAKES ENTOMOLOGIST 265 Regarding host data, Stoner (1922) and McPherson (1970, 1979b) provided a list of host plants, often from extra-Michigan sources. However, I have not reproduced those lists of plants provided by Stoner, McPherson, and other authors but essentially limited this treatment to those data from Michigan-collected specimens. Collections are designated as follows: Daniel R. Swanson, personal collection (DRS); Albert J. Cook Arthropod Research Collection, Michigan State University, East Lansing, Michigan (MSUC); and University of Michigan Mu- seum of Zoology Insect Collection, Ann Arbor, Michigan (UMMZ).

Results and Discussion Superfamily PENTATOMOIDEA The Pentatomoidea are a superfamily of true bugs comprising 352 species in 103 genera in 7 families in the United States. Treated as the Scutelleroidea in the past, they are informally recognized by the usually 5-segmented antennae and enlarged scutellum, which completely covers the body in some groups; these features, combined with an ovoid or elliptical shape, make members of this group generally easy to recognize. The paired trichobothria (single in the Podopinae) of the abdominal sternites also is an important distinguishing character (Henry 1997). Most are uni- or bivoltine, overwinter as adults, and lay eggs in the spring (McPherson 1982). Additionally, many species are well- known and frequently encountered, as pests of gardens and commercial crops, singly or in large aggregations around lights at night (i.e., McPherson and Sites 1989), or as conspicuous overwintering guests in human domiciles. Most are phytophagous, often feeding on a wide range of plants, although the members of one subfamily (Pentatomidae: Asopinae) are predaceous. The group contains many economically important taxa; references relating to particular groups or species are mentioned under their respective headings. The acoustic behavior viz. stridulation of members of this group was summarized by Schaefer (1980). The genitalia often provide the only dependable character for unequivocally identifying species, and those of selected species were described by McDonald (1966) and Davidová-Vilímová and McPherson (1991); these also are mentioned under the species accounts, where applicable. Six chapters by Froeschner (1988a, 1988b, 1988c, 1988d, 1988e, 1988f) compose the most current catalog for the taxa found in America north of Mexico. Blatchley (1926) and Torre-Bueno (1939) presented keys to the species of the eastern and continental United States, respectively, but out-dated and erroneous data diminish their utility. Rolston and McDonald (1979) and Schuh and Slater (1995) discussed the relationships of and provided keys for the world families of the Pentatomoidea, and Grazia et al. (2008) provided a cladistic analysis for the superfamily. Many of the species found in Michigan are treated by Hoffman (1971); yet, McPherson’s (1982) seminal treatment of the Northeastern Pentatomoidea remains one of the most complete and informative. Five of the seven subfamilies found in the Nearctic region are represented in Michigan; the Plataspidae and Tessaratomidae, each represented by a single species, have a more southerly distribution. Seventy-five species in 43 genera are found in the state (Table 1). The key presented below is synthesized primarily from works by McPher- son (1970, 1982), Rolston and McDonald (1979), and Schuh and Slater (1995). The difficulty created in the key by the distribution of the greatly enlarged scutellum within the group is worth noting, and other characters might provide a “cleaner” criterion for identification. The natural relationships, however, become even more greatly obscured in several other iterations; thus, I have opted for the form presented below. 266 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Table 1. List of the Michigan Pentatomoidea.

Scutelleridae Rhacognathus americanus (Stål), 1870 Acantholomidea denticulata (Stål), 1870 Stiretrus anchorago (Fabricius), 1775 Eurygaster alternata (Say), 1828 Zicrona caerulea (Linnaeus), 1758 Homaemus aeneifrons (Say), 1824 Phimodera binotata (Say), 1824 Pentatomidae: Pentatominae Tetyra bipunctata (Herrich-Schaeffer), 1839 Aelia americana Dallas, 1851 Banasa calva (Say), 1832 Thyreocoridae Banasa dimidiata (Say), 1832 Corimelaena agrella McAtee, 1919 Banasa sordida (Uhler), 1871 Corimelaena lateralis lateralis (Fabricius), Brochymena quadripustulata (Fabricius), 1803 1775 Corimelaena nigra Dallas, 1851 Chinavia hilaris (Say), 1832 Corimelaena obscura McPherson and Sailer, Chinavia pensylvanica (Gmelin), 1790 1978 Chlorochroa persimilis Horvath, 1908 Corimelaena pulicaria (Germar), 1839 Coenus delius (Say), 1832 Cydnoides ciliatus orientis McAtee and Cosmopepla lintneriana Kirkaldy, 1909 Malloch, 1933 Dendrocoris humeralis (Uhler), 1877 Galgupha aterrima Malloch, 1919 Euschistus ictericus (Linnaeus), 1763 Galgupha atra Amyot and Serville, 1843 Euschistus politus Uhler, 1897 Galgupha carinata McAtee and Malloch, Euschistus servus euschistoides 1933 (Vollenhoven), 1868 Galgupha loboprostethia Sailer, 1940 Euschistus servus servus (Say), 1832 Galgupha nitiduloides nitiduloides (Wolff), Euschistus tristigmus luridus Dallas, 1851 1802 Euschistus tristigmus tristigmus (Say), 1832 Galgupha ovalis Hussey, 1925 Euschistus variolarius (Palisot de Beauvois), 1817 Cydnidae Halyomorpha halys (Stål), 1855 Amnestus pallidus Zimmer, 1910 Holcostethus fulvipes (Ruckes), 1957 Amnestus pusillus Uhler, 1876 Holcostethus limbolarius (Stål), 1872 Amnestus spinifrons (Say), 1825 Holcostethus mcdonaldi Rider and Rolston, Melanaethus robustus Uhler, 1877 1995 Pangaeus bilineatus (Say), 1825 Hymenarcys nervosa (Say), 1832 Sehirus cinctus albomaculatus Dallas, 1851 Mcphersonarcys aequalis (Say), 1832 Sehirus cinctus cinctus (Palisot de Beau- Menecles insertus (Say), 1832 vois), 1811 Mormidea lugens (Fabricius), 1775 Neottiglossa trilineata (Kirby), 1837 Acanthosomatidae Neottiglossa undata (Say), 1832 Elasmostethus atricornis (Van Duzee), 1904 Oebalus pugnax (Fabricius), 1775 Elasmostethus cruciatus (Say), 1831 Parabrochymena arborea (Say), 1825 Elasmucha lateralis (Say), 1831 Prionosoma podopioides Uhler, 1863 Sciocoris microphthalmus Flor, 1860 Pentatomidae: Asopinae Thyanta calceata (Say), 1832 Apoecilus bracteatus (Fitch), 1856 Thyanta custator accerra McAtee, 1919 Apoecilus cynicus (Say), 1832 Trichopepla atricornis (Stål), 1872 Perillus bioculatus (Fabricius), 1775 Trichopepla semivittata (Say), 1832 Perillus circumcinctus (Stål), 1862 Perillus exaptus (Say), 1825 Pentatomidae: Podopinae Podisus brevispinus Phillips, 1982 Amaurochrous brevitylus Barber and Sailer, Podisus maculiventris (Say), 1832 1953 Podisus neglectus (Westwood), 1837 Amaurochrous cinctipes (Say), 1828 Podisus placidus Uhler, 1870 Podisus serieventris Uhler, 1871

2012 THE GREAT LAKES ENTOMOLOGIST 267 Key to the North American families of Pentatomoidea 1 Tibiae armed with numerous strong spines; [tarsi 3-segmented]...... 2 1’ Tibiae not armed with numerous strong spines...... 3 2 (1) Scutellum greatly enlarged, U-shaped, covering most of hemelytra and abdomen...... Thyreocoridae 2’ Scutellum not greatly enlarged, more or less triangular, leaving hemelytra and abdomen broadly exposed...... Cydnidae 3 (1’) Scutellum greatly enlarged, U-shaped, covering most of hemelytra and abdomen...... 4 3’ Scutellum not greatly enlarged, more or less triangular (except Stire- trus (Pentatomidae: Asopinae), then with bright contrasting colors), leaving hemelytra and abdomen broadly exposed (except Podopinae (Pentatomidae), then with prominent humeral projection)...... 5 4 (3) Hemelytra much longer than abdomen, bent between corium and membrane and folded below scutellum in repose; abdominal sterna with straight, transverse sulcus on each side; tarsi 2-segmented [not present in Michigan]...... Plataspidae 4’ Hemelytra, at most, only slightly longer than abdomen, not bent at junction of corium and membrane; abdominal sterna lacking transverse sulci; tarsi 3-segmented...... Scutelleridae 5 (3’) Pronotum extending over base of scutellum; spiracles of second abdominal sternite exposed, not hidden by the metapleura, a short but evident distance from posterior margin; [tarsi 3-segmented] [not present in Michigan]...... Tessaratomidae 5’ Pronotum ending at base of scutellum; spiracle of second abdominal sternite concealed by metapleura, rarely exposed just at the posterior margin of metasternum...... 6 6 (5’) Tarsi 2-segmented; thoracic and abdominal sterna with distinct midlongitudinal carina...... Acanthosomatidae 6’ Tarsi 3-segmented; thoracic and abdominal sterna without distinct longitudinal carina...... Pentatomidae

Family SCUTELLERIDAE Leach, 1815 Commonly known as shield bugs, the Scutelleridae are recognized by the greatly enlarged, rounded scutellum, which covers most of the abdomen (although this feature is present in a few other groups, most noticeably the Thyreocoridae). Its members are marked by great variation in color and color pattern such that these characters rarely can be used in species recognition. The group has been accorded variously familial or subfamilial rank under the Pentatomidae, although contemporary treatments almost unanimously assign the higher status. Javahery et al. (2000) discussed the economic importance of this group, and some Old World members of the group may cause severe agricultural damage in large numbers. Lattin (1964) provided the most recent comprehensive treatment of the U.S. species; unfortunately, as an unpublished dissertation, its taxonomic acts are not recognized as valid. I have chosen to include information that relates to the Michigan taxa, although the aforementioned admonition should be heeded. Lattin’s (1964) county records are included, where ascertain- able by the author, without reservation, as repositories or ownership were included with the specimen data; these few instances are noted under the appropriate entry. 268 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

In the United States, the Scutelleridae comprise 37 species in 16 genera (Froeschner 1988d). Members of all four New World subfamilies may be found north of Mexico (Hoteinae contains only Old World taxa), and three of those subfamilies are represented in Michigan; members of the nominate Scutellerinae do not occur in the state. Five genera are each represented by a single species in Michigan. The provided key was modified from McPherson’s (1970) treatment, with the subfamily framework adapted from Schuh and Slater (1995).

Key to the Scutelleridae of Michigan 1 All abdominal sterna lacking stridulatory areas...... 2 1’ Stridulatory area present on fourth, fifth, and sometimes sixth abdominal sternite (Pachycorinae)...... 3 2 (1) Scutellum narrower, exposing hemelytra for entire length and broadly exposing connexiva; wings with intervannal vein well developed; ostiole distinct, with evident canal (Eurygastrinae)...... Eurygaster alternata 2’ Scutellum broadly developed, exposing hemelytra only near base and only narrowly exposing connexiva; wing with intervannal vein greatly reduced; ostiole indistinct, without evident canal (Odontotarsinae)...... Phimodera binotata 3 (1’) Pronotum with distinct transverse groove; head distinctly deflexed; lateral margins of head and pronotum finely and irregularly dentate...... Acantholomidea denticulata 3’ Pronotum without transverse groove; head gradually declivent; lateral margins of head and pronotum entire...... 4 4 (3’) Ostiolar opening not extended as canal towards lateral margin of metapleuron; length 12 mm or more...... Tetyra bipunctata 4’ Ostiolar opening extended as long, slender canal towards lateral margin of metapleuron; length 9 mm or less...... Homaemus aeneifrons

Subfamily EURYGASTRINAE Amyot & Serville, 1843 Genus EURYGASTER Laporte, 1832 Eurygaster alternata (Say), 1828. – This species was reported from Michi- gan by Stoner (1922), who frequently found it “in open, dry grassy situations... particularly on the higher ground.” Hussey (1922a), however, described E. alternata as “[m]oderately common in marshy localities, where it occurs among the grasses and sedges”, suggesting this scutellerid may thrive in habitats of varying wetness. Label data indicate E. alternata has been collected from fruit- ing sedge heads in Livingston County and a window pane trap in Montmorency County. This species also has been collected on Isle Royale (Keweenaw County). Javahery et al. (2000) discussed the economic importance of several species in this genus. McDonald (1966) described the genitalia. Vojdani (1961) reviewed the species of Eurygaster found in the United States. 128 specimens examined. Collection dates from 14 May to 13 September. Distribution: Alger, Allegan, Alpena, Barry, Berrien, Charlevoix, Cheboy- gan, Chippewa, Clare, Crawford, Delta, Dickinson, Emmet, Gogebic, Grand Tra- verse, Houghton, Huron, Iosco, Kent, Keweenaw, Lake, Livingston, Mackinac, Marquette, Mecosta, Menominee, Midland, Montcalm, Montmorency, Muskegon, Newaygo, Oscoda, Otsego, Schoolcraft, Van Buren, and Washtenaw counties. Eurygaster amerinda Bliven, 1956. – Lattin (1964) reported his new subspecies, Eurygaster amerinda knighti, from Michigan, although it was not included in the state faunal list by McPherson (1970), perhaps for the reasons associated 2012 THE GREAT LAKES ENTOMOLOGIST 269 with doctoral dissertations noted above. Subsequently, McPherson (1980a) did include the reference in his treatment of the northeastern states. Lattin’s maps include records for this species in what appear to be Cheboygan and Marquette counties as well as Isle Royale (Keweenaw County). The author has not examined any specimens that might be unequivocally referred to this species.

Subfamily ODONTOTARSINAE Stål, 1872 Genus PHIMODERA Germar, 1839 Phimodera binotata (Say), 1824. – Lattin (1964) and McPherson (1970) recorded this species from Michigan, and the former author’s record for Che- boygan County is included in the distribution list. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Reuter (1906) monographed the genus (in German), although P. binotata is treated only peripherally in that work. 3 specimens examined. Collection dates from 24 July to 18 August. Distribution: Charlevoix, Cheboygan, and Mackinac counties. Phimodera torpida Walker, 1867. – This species was reported from Michi- gan by Steyskal (1938), and the record was included by Froeschner (1988d). Steyskal included the locality data, and the specimen on which this record is based resides in the UMMZ. However, Hussey modified his identification since Steyskal’s publication as indicated by the 1921 determination label for P. torpida being folded up and re-placed on the pin, with a new determination label for P. binotata being added in 1956. Lattin (1964) synonymized this (and another) species with P. binotata, negating this issue had the act been considered valid. This specimen was included in McPherson’s (1970) treatment under P. binotata. Thus, P. torpida, an otherwise western species recorded from Colorado, Utah, Alberta, British Columbia, and Saskatchewan (Froeschner 1988d), is not known from Michigan and should be excluded from the faunal list.

Subfamily PACHYCORINAE Amyot & Serville, 1843 Genus ACANTHOLOMIDEA Sailer, 1945 Acantholomidea denticulata (Stål), 1870. – Lattin (1964) and McPherson (1970) recorded this species from Michigan, and the former author’s record for Cheboygan County is included in the distribution list. The dentate margin of the pronotum is unique among the Michigan scutellerids. Harris and Andre (1934) reported biological notes on A. denticulata, including its occurrence on Ceanothus pubescens (Watson) and C. ovatus Desfontaines. 2 specimens examined. Collection dates from 6 May to 7 July. Distribution: Berrien, Cheboygan, Saint Joseph, and Washtenaw counties.

Genus HOMAEMUS Dallas, 1851 Homaemus aeneifrons (Say), 1824. – This species was reported from Michi- gan by Stoner (1922), who noted it fairly common in dry habitats (i.e., sandy or grassy high ground) as well as wet ones (i.e., bogs and creek sides). This species has been collected from flowers ofRubus sp. (Rosaceae) in Iron County and “[s] weeping brackensweet fern fallow field” in Isabella County. It also has been collected on Isle Royale (Keweenaw County). The shiny black head may serve as a visual cue in separating this species from E. alternata, a species which, while not closely related, resembles H. aeneifrons in color and form. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Walley (1929) keyed the species of Homaemus found north of Mexico. 344 specimens examined. Collection dates from 26 June to 16 October. 270 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Distribution: Alcona, Alger, Alpena, Baraga, Barry, Benzie, Charlevoix, Cheboygan, Chippewa, Clare, Crawford, Delta, Dickinson, Emmet, Gogebic, Grand Traverse, Houghton, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalkas- ka, Keweenaw, Lenawee, Livingston, Luce, Mackinac, Marquette, Menominee, Midland, Missaukee, Montmorency, Oakland, Ogemaw, Ontonagon, Oscoda, Otsego, Presque Isle, Roscommon, and Schoolcraft counties.

Genus TETYRA Fabricius, 1803 Tetyra bipunctata (Herrich-Schaeffer), 1839. – Lattin (1964) and subsequently McPherson (1970) reported the shield-backed pine seed bug from Michigan, with each noting its proclivity for attacking pines. Label data from examined specimens corroborate this predilection, indicating its collection from jack pine (Pinus banksiana Lamb.) and European black pine (Pinus nigra Arnold) in Saint Joseph County. Gilbert et al. (1967) treated the bionomics of T. bipunctata in association with jack pine in Wisconsin, and Javahery et al. (2000) discussed the economic importance of this species. Additional label data indicate several individuals were taken on the Lake Michigan shoreline in Manistee County. The large size of this scutellerid will separate it instantly from all other members of the family in Michigan. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. 32 specimens examined. Collection dates from 26 May to 18 September. Distribution: Cass, Cheboygan, Grand Traverse, Iosco, Leelanau, Man- istee, Marquette, Montmorency, Oscoda, and Saint Joseph counties.

Family THYREOCORIDAE Amyot & Serville, 1843 Commonly referred to as the ebony or negro bugs, the small shiny black members of this group are chiefly recognized by the enlarged scutellum which covers most of the abdomen and wings, a condition also found in the Scutelleridae. The scutellum, along with the small black habitus, render these insects frequently confused with beetles. Thyreocorids are frequently found on short vegetation, feeding on flowers and developing seeds. The group is not generally regarded to be of economic importance (Lis et al. 2000), although some of its members negatively affect the taste of berries on which they feed (Metcalf et al. 1962). The taxonomic position of the Thyreocoridae has varied greatly, as full familial status or as subfamilies of the Pentatomidae or Cydnidae; several contemporary treatments (i.e., Dolling 1981, Schuh and Slater 1995, Lis et al. 2000) relegate them to subfamilial status under the latter. Here the more traditional treatment is followed, without intentional support of either position. There has also been contention regarding the family name (i.e., Rolston and McDonald 1979, Štys and Davidová 1979), and although many authors have used Corime- laenidae to refer to the group in the past, I have used the Thyreocoridae here in keeping with the current catalog (Froeschner 1988f). Ahmad and McPherson (1990) described the male genitalia and its taxonomic significance for species in each of the three genera here included. The Thyreocoridae comprises 41 species in 4 genera in the United States (Froeschner 1988f). McAtee and Malloch (1933) treated the group (as a subfamily of the Pentatomidae), and this remains perhaps the most relevant for the taxa of the United States. All species found north of Mexico belong to the subfamily Corimelaeninae. Twelve species in 3 genera are known from Michigan, and the key provided is modified from McPherson’s (1982) treatment of the northeastern Pentatomoidea. 2012 THE GREAT LAKES ENTOMOLOGIST 271 Key to the Thyreocoridae of Michigan 1 Lateral margins of pronotum and abdomen ciliate; [costal margin bordered within by definite groove; spiracles nearer to trichobothria than to lateral margins of abdominal sternites; corium entirely black, never with pale costal margin]...... Cydnoides ciliatus orientis 1’ Lateral margins of body not ciliate...... 2 2 (1’) Pronotum and scutellum, in lateral view, not forming continuous convex line but with distinct transverse declivity between; costal margin of hemelytra not bordered within by groove; spiracles nearer to lateral margins of abdominal sternites than to trichobothria (Corimelaena)...... 3 2’ Pronotum and scutellum, in lateral view, forming continuous convex line; costal margin of hemelytra bordered within by definite groove; spiracles nearer to trichobothria than to lateral margins of abdominal sternites; [corium entirely black, never with pale costal margin] (Gal- gupha)...... 7 3 (2) Apex of corium acute; [corium with pale costal margin, inner margin of pale costal stripe slightly angulate near middle; length greater than 3.5 mm]...... Corimelaena agrella 3’ Apex of corium rounded or nearly so...... 4 4 (3’) Corium entirely black, distinctly punctured to costal margin; [length greater than 3.5 mm]...... Corimelaena nigra 4’ Corium with pale costal margin, almost impunctate along costa...... 5 5 (4’) Pale costal stripe widened basally, extending over cubital vein; size smaller, length 3.5 mm or less...... Corimelaena pulicaria 5’ Pale costal stripe narrowed basally, not extending over cubital vein; size larger, length usually greater than 3.5 mm...... 6 6 (5’) Pale costal stripe well-defined, entire; first conjunctival appendages usually serrate only on outer margins (occasionally with 1-2 teeth on inner margins) with apical hooks obtusely rounded; genital plates usually subtriangular...... Corimelaena lateralis lateralis 6’ Pale costal stripe invaded and frequently almost obscured by fuscous coloration, particularly in middle third; first conjunctival appendages strongly serrate on inner and outer margins with gradually tapering and acute apical hooks; genital plates usually subquadrangular...... Corimelaena obscura 7 (2’) Metapleuron with lateral area adjacent to evaporatorium distinctly punctate (subgenus Nothocoris)...... Galgupha nitiduloides nitiduloides 7’ Metapleuron with lateral area impunctate or nearly so (subgenus Galgupha)...... 8 8 (7’) Prostethium with anterior margin produced to form explanate lobe, granulate portion as wide as base of second rostral segment...... Galgupha loboprostethia 8’ Prostethium with anterior margin not lobed, granulated portion narrow...... 9 9 (8’) Anterodorsal series of spines on each protibia with distal two spines either decidedly weak or absent; corium without distinct ridge inside costal groove, ridge represented only by series of punctures distally; punctation on discal portions of pronotum and scutellum mostly obsolete...... 10 9’ Anterodorsal series of spines on each protibia terminating in two spines equal or subequal in size to remaining spines in series; corium 272 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

with distinct ridge inside costal groove; punctation on discal portions of pronotum and scutellum more evident...... 11 10 (9) Outline, in dorsal view, distinctly more narrowly rounded posteriorly than anteriorly; scutellum thickened along apical margin; outline of pygophore, in dorsal view, subquadrate, dorsal rim with well-developed carinate elevation on each inner side anteriorly; genital plates about two-thirds the length of sixth abdominal sternite...... Galgupha carinata 10’ Outline, in dorsal view, usually scarcely more narrowly rounded posteriorly than anteriorly; scutellum not thickened along apical margin; outline of pygophore, in dorsal view, subtriangular, broadly basin-like giving cup a scooped out appearance, dorsal rim without anterior carinae; genital plates about half the length of sixth abdominal sternite...... Galgupha atra 11 (9’) Scutellum abruptly declivent posteriorly; sixth sternite angulate in both sexes; branches of exocorial vein subparallel; posterior border of pygophore, in ventral view, strongly concave, dorsal rim with numerous long hairs posteriorly...... Galgupha aterrima 11’ Scutellum gradually declivent posteriorly; sixth sternite rounded to subangulate in both sexes; branches of exocorial vein noticeably diver- gent; posterior border of pygophore, in ventral view, weakly concave, dorsal rim without numerous long hairs posteriorly...... Galgupha ovalis

Subfamily CORIMELAENINAE Uhler, 1871 Genus CORIMELAENA White, 1839 Corimelaena agrella McAtee, 1919. – (NEW STATE RECORD). A single specimen of this thyreocorid recently was collected by the author. Label data as follows: MICHIGAN: Jackson Co., Sharonville State Wildlife Mngmt. Area, swept from open grassy field, 27 May 2012, 42.1875°N 84.1443°W, 990 ft., D. R. Swanson, #33, det. D. R. Swanson 2012 [1 male] (DRS). This species is otherwise known from Iowa, Illinois, Kentucky, Maryland, Texas, and Washing- ton, D.C. (Froeschner 1988f). Although its recorded range might not strongly support a Michigan extension, the specimen undoubtedly belongs to a species heretofore unknown from Michigan as it does not possess a rounded corial apex. Furthermore, the larger size and slight inner angulation of the marginal corial stripe (figured by McPherson (1982)) leaves little doubt to its specific identity. 1 specimen examined. Collection date is 27 May. Distribution: Jackson County. Corimelaena lateralis lateralis (Fabricius), 1803. – This species was reported from Michigan by Townsend (1890) where it was commonly taken on flowers. Label data indicate C. l. lateralis has been collected from golden alexander (Zizia aurea (L.) W. D. J. Koch) in Washtenaw County. McPherson (1971a, 1972a) described the life history and rearing of this species on wild carrot. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. 118 specimens examined. Collection dates from 14 May to 4 October. Distribution: Bay, Berrien, Calhoun, Cass, Clinton, Gladwin, Gratiot, Hill- sdale, Huron, Ingham, Ionia, Isabella, Jackson, Kalamazoo, Lapeer, Lenawee, Livingston, Macomb, Menominee, Midland, Monroe, Montcalm, Montmorency, Oakland, Saint Joseph, Shiawassee, Tuscola, Van Buren, Washtenaw, and Wayne counties. Corimelaena nigra Dallas, 1851. – McAtee and Malloch (1933) reported this species from Michigan. Label data indicate C. nigra has been collected from a black light in Ottawa County and a Malaise trap in Lake County. It also 2012 THE GREAT LAKES ENTOMOLOGIST 273 has been collected on Isle Royale (Keweenaw County). 23 specimens examined. Collection dates from 26 April to 10 August. Distribution: Barry, Berrien, Cheboygan, Chippewa, Clinton, Kalamazoo, Keweenaw, Lake, Lapeer, Livingston, Luce, Mackinac, Marquette, Midland, Muskegon, Oakland, Ottawa, Presque Isle, and Washtenaw counties. Corimelaena obscura McPherson and Sailer, 1978. – This species was reported from Michigan in the original description (McPherson and Sailer 1978) based on a single specimen collected in Wayne County on 2 April; McPherson (1979b) subsequently mentioned the record. Bundy and McPherson (1997) described the life history and laboratory rearing of this species. 1 specimen examined. Collection dates from 2 April to 21 June. Distribution: Clinton and Wayne counties. Corimelaena pulicaria (Germar), 1839. – A common thyreocorid in Michi- gan, C. pulicaria was first recorded from the state by Townsend (1890) who noted it frequently taken on flowers. Stoner (1922) took the species in a field with redtop (Agrostis sp.), timothy (Phleum pratense L.), and bristly crowfoot (Ranunculus pennsylvanicus L.f.). Hussey (1922a) found it “taken from ragweed (Ambrosia) and a few were found in marsh grasses…several were collected from roadside grasses.” Label data indicate the species has been swept from open fields in Oakland County and lakeplain prairies in Allegan and Saint Clair counties as well as collected on beaches in Berrien County. One collection in Isabella County was described as “sweeping fallow roadside. Solidago dominant. Some shrubs.” Further label data also indicate some plants from which this species was collected: willow (Salix sp.) in Huron County, alfalfa (Medicago sativa L.) in Ingham County, and gray goldenrod (Solidago nemoralis Aiton) in Livingston County. This insect has been collected from a rotary trap in a field of oats in Berrien County and a pitfall trap in Ingham County. It is well documented that secretions given off by this insect may unpleasantly alter the taste of berries on which it feeds (Metcalf et al. 1962). McDonald (1966) described the genitalia. 425 specimens examined. Collection dates from 8 April to 13 November. Distribution: Alcona, Allegan, Arenac, Barry, Bay, Berrien, Branch, Cass, Charlevoix, Cheboygan, Chippewa, Clinton, Crawford, Dickinson, Eaton, Em- met, Genesee, Gladwin, Gogebic, Grand Traverse, Gratiot, Hillsdale, Houghton, Huron, Ingham, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Livingston, Luce, Mason, Mecosta, Midland, Missaukee, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Ontonagon, Otsego, Presque Isle, Roscommon, Saginaw, Saint Clair, Saint Joseph, Shiawassee, Tuscola, Van Buren, Washtenaw, and Wayne counties.

Genus CYDNOIDES Malloch, 1919 Cydnoides ciliatus orientis McAtee and Malloch, 1933. – This species was reported from Michigan by McPherson (1979b). Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. 2 specimens examined. Collection dates from 13-17 August. Distribution: Van Buren County.

Genus GALGUPHA Amyot & Serville, 1843 Subgenus GALGUPHA Amyot & Serville, 1843 Galgupha aterrima Malloch, 1919. – This species was reported from Michigan by Hussey (1922a) where “[o]ne was swept from ground vegetation on the dunes.” This species has been taken from a rotary trap in an oat field and a wheat field in Berrien and Kalamazoo County, respectively, as well as from a pyramid trap in Leelanau County. McPherson (1978c) re-examined type 274 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 material of this species. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. 69 specimens examined. Collection dates from 22 March to 13 November. Distribution: Berrien, Charlevoix, Cheboygan, Crawford, Huron, Ingham, Kalamazoo, Kent, Lapeer, Leelanau, Mason, Montmorency, Muskegon, Saint Joseph, Van Buren, Washtenaw, Wayne, and Wexford counties. Galgupha atra Amyot and Serville, 1843. – This species was reported from Michigan by Townsend (1890) (as Corimelaena atra) where it was commonly taken on flowers. Stoner (1922) collected purported nymphs of this species from a low grassy creekbed and found several adults in a low field of redtop (Agrostis sp.); he also recorded it (as Thyreocoris ater) from beach drift. Hussey (1922a) noted them “swept from a tangle of grasses and vines on the flood plain of the Galien River.” Label data indicate this species has been taken from a Malaise trap in Monroe County and on Virginia strawberry (Fragaria virginiana Duch- esne) in Livingston County. 58 specimens examined. Collection dates from 13 May to 12 September. Distribution: Allegan, Barry, Benzie, Berrien, Cass, Charlevoix, Che- boygan, Clare, Clinton, Crawford, Dickinson, Emmet, Gogebic, Kalamazoo, Kalkaska, Livingston, Mackinac, Macomb, Marquette, Mecosta, Midland, Monroe, Oakland, Ogemaw, Saginaw, Saint Joseph, Schoolcraft, Van Buren, Washtenaw, and Wexford counties. Galgupha carinata McAtee and Malloch, 1933. – This species was reported from Michigan by McPherson (1970). 3 specimens examined. Collection date is 2 August. Distribution: Washtenaw County. Galgupha loboprostethia Sailer, 1940. – This species was reported from Michigan by McPherson (1970). This species has been collected from a Malaise trap in Berrien County. Sailer’s (1940) original description and subsequent notes (Sailer 1941) constitute the bulk of the information on this purportedly rare species. 6 specimens examined. Collection dates from 22 May to 8 August. Distribution: Allegan, Berrien, Jackson, Mason, and Otsego counties. Galgupha ovalis Hussey, 1925. – This species was reported from Michigan by McPherson (1970). Biehler and McPherson (1982) described the life history and laboratory rearing of this thyreocorid. 19 specimens examined. Collection dates from 27 April to 13 November. Distribution: Allegan, Benzie, Berrien, Cheboygan, Clare, Ingham, Ionia, Livingston, and Oceana counties.

Subgenus NOTHOCORIS McAtee and Malloch, 1928 Galgupha nitiduloides nitiduloides (Wolff), 1802. – This species was reported from Michigan by Townsend (1890) (as Corimelaena nitiduloides) “taken in the rubbish on the top of an ant-hill belonging to Formica schaufussi [Mayr, 1866]. This species is commonly…found on flowers.” Subsequently, Stoner (1922) and Hussey (1922a) each recorded it from Michigan; the latter noted them “taken from roadside grasses…and another was beaten from vegetation in a clearing behind the Sawyer Dunes.” Label data indicate G. n. nitiduloides has been taken under debris on the Lake Michigan shore in Allegan County. It also has been collected on Isle Royale (Keweenaw County). McDonald (1966) described the female genitalia. 26 specimens examined. Collection dates from 22 May to 13 November. Distribution: Allegan, Barry, Berrien, Cass, Cheboygan, Clinton, Delta, Dickinson, Houghton, Ingham, Kalamazoo, Kalkaska, Keweenaw, Mason, Mid- land, Missaukee, Montcalm, Newaygo, Roscommon, Saint Joseph, Schoolcraft, and Wayne counties. 2012 THE GREAT LAKES ENTOMOLOGIST 275 Family CYDNIDAE Billberg, 1820 The burrowing (or burrower) bugs, aptly named for their hypogean life- style, are typically small and black, like the thyreocorids, but lack the distinct enlarged scutellum found in that family. Cydnids generally feed on roots and other subterranean plant material (except perhaps the Sehirinae), and it has been suggested that the members of this family are univoltine, lay eggs in or on the soil, and may be attracted to lights during the surface-dwelling portion of their life (Froeschner 1960). These insects are not thought to be particularly injurious to those plants of agricultural significance to mankind, and Lis et al. (2000) summarized the economic importance of the group. Members of this family also have had a high tendency for accidental introduction through soil ballasts in old sailing ships (Froeschner 1960). The position of the Cydnidae, like many other pentatomoid family-groups, has experienced several changes. A wide view of the family (ie. Dolling 1981, Schuh and Slater 1995) subsumes the Thyreocoridae within it, but the more traditional view is followed here (as mentioned under the Thyreocoridae). The Cydnidae also has been accorded subfamilial rank under the Pentatomidae in the past. The Cydnidae comprises 43 species in 13 genera in the United States (Froeschner 1988b). Four of six world subfamilies may be found north of Mexico, but only three of those are represented in Michigan; species of the Scaptocorinae are known only from the southeastern states. Froeschner’s (1960) monograph remains the most important reference for the taxa north of Mexico. Six species in 4 genera are known from Michigan, and the two new state records presented below encompass the first record of the nominate subfamily, Cydninae, for Michigan. The following key was modified from Froeschner’s (1960) treatment.

Key to the Cydnidae of Michigan 1 Clavi meeting beyond scutellum and forming commissure; anterior margin of head with distinct peg-like teeth (Amnestinae: Amnestus)...... 2 1’ Clavi not meeting beyond scutellum, not forming claval commissure; anterior margin of head without distinct peg-like teeth...... 4 2 (1) Jugum with four marginal pegs; smaller species, length usually less than 2.5 mm; [rostrum attaining base of metacoxae]...... Amnestus pusillus 2’ Jugum with five or more marginal pegs; larger species, length usually greater than 2.5 mm...... 3 3 (2’) Rostrum not reaching base of abdomen, attaining, at most, middle of metasternum; color usually reddish-brown...... Amnestus pallidus 3’ Rostrum long, at least reaching base of abdomen; color usually dark chestnut brown...... Amnestus spinifrons 4 (1’) Pronotum without lateral, submarginal row of setigerous punctures; second tarsomere distinctly narrower than first and third (Sehirinae: Sehirus cinctus)...... 5 4’ Pronotum with lateral, submarginal row of setigerous punctures; all tarsomeres subequal in diameter (Cydninae)...... 6 5 (4) Corium marked with white spot at apex of radial vein; jugal margins elevated above dorsum of head in profile...... Sehirus cinctus albomaculatus 5’ Corium not marked with white at apex of radial vein; jugal margins lower than dorsum of head in profile...... Sehirus cinctus cinctus 276 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

6 (4’) Anterior part of osteolar peritreme modified apically into distinct simple, flat, posteriorly differentiated polished lobe which is wider than basal part of peritreme (Melanaethus); pronotum anteriorly without distinctly impressed line; jugum with one submarginal seta; size smaller, length less than 5 mm...... Melanaethus robustus 6’ Anterior part of osteolar peritreme without differentiated apical structure; pronotum anteriorly with deep, sharply impressed impunctate line paralleling anterior margin from side to side; jugum with three or more submarginal setae; size larger, length greater than 5 mm...... Pangaeus bilineatus

Subfamily AMNESTINAE Hart, 1919 Genus AMNESTUS Dallas, 1851 Amnestus pallidus Zimmer, 1910. – Froeschner (1960) reported this species from Michigan. Label data indicate A. pallidus has been taken from white and ultraviolet lights in Clinton County. It also has been collected from pitfall and window pane traps in Ingham and Kalkaska County, respectively. McDonald (1966) described the genitalia. 28 specimens examined. Collection dates from 26 April to 9 August. Distribution: Clinton, Ingham, Kalkaska, Midland, Saginaw, and Washt- enaw counties. Amnestus pusillus Uhler, 1876. – This species was reported from Michigan by McPherson (1970). Label data indicate it has been collected in a window pane trap in Kalkaska County. 39 specimens examined. Collection dates from 8 May to 5 September. Distribution: Alpena, Cheboygan, Clare, Clinton, Gladwin, Ingham, Iosco, Kalamazoo, Kalkaska, Lenawee, Livingston, Midland, Monroe, Osceola, Roscom- mon, and Washtenaw counties. Amnestus spinifrons (Say), 1825. – Parshley (1917) reported this species from Michigan. Label data indicate A. spinifrons has been taken on red clover (Trifolium pratense L.) in Kalamazoo County and from a rotary trap in a field of oats and on a beach in Berrien County. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. 14 specimens examined. Collection dates from 19 April to October [no date]. Distribution: Berrien, Clinton, Ingham, Kalamazoo, Kent, Lenawee, Ma- son, Midland, and Washtenaw counties.

Subfamily CYDNINAE Billberg, 1820 Genus MELANAETHUS Uhler, 1876 Melanaethus robustus Uhler, 1877. – (NEW STATE RECORD). Three specimens of this small cydnine were collected by the author in the late spring of 2012. Label data as follows: MICHIGAN: Oakland Co., Highland, Highland State Rec. Area, swept from low grass in open woods, 20 May 2012, 42.6427°N 83.5536°W, 870 ft., D. R. Swanson, #29, det. D. R. Swanson 2012 [1 female] (DRS); MICHIGAN: Jackson Co., Sharonville State Wildlife Mngmt. Area, swept from open grassy field, 27 May 2012, 42.1875°N 84.1443°W, 990 ft., D. R. Swanson, #33, det. D. R. Swanson 2012 [2 females] (DRS). The occurrence of this species in Michigan is not implausible as it is known from Illinois, In- diana, and Ohio, among other states (Froeschner 1988b). The Jackson County record is approximately 225 air kilometers northwest of Delaware County, Ohio and 280 air kilometers west of Ashtabula County, Ohio, the nearest localities reported by Froeschner (1960). Davidová-Vilímová and McPherson 2012 THE GREAT LAKES ENTOMOLOGIST 277 (1991) described and illustrated the male genitalia. 3 specimens examined. Collection dates from 20 to 27 May. Distribution: Jackson and Oakland counties.

Genus PANGAEUS Stål, 1862 Pangaeus bilineatus (Say), 1825. – (NEW STATE RECORD). To the author’s knowledge, this species has never been reported from Michigan. This fact, perhaps partially explained by the lack of representation in the collections (all but three specimens came from the author’s personal collection), is surprising considering the larger habitus and relative abundance of this species, especially given the depauperate cydnid fauna of Michigan. Label data indicate P. bilineatus has been collected from white and ultraviolet lights in Clinton County. In Washtenaw County, the author has taken several individuals crawling on concrete sidewalks and parking lots on sunny days in early summer. There has been some question as to whether this cydnid might adversely affect growth of spinach and peanut crops (Gould 1931, Smith and Pitts 1974, Chapin and Thomas 2003, Chapin et al. 2006), and Lis et al. (2000) summarized the literature regarding the economic importance of this species. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. Sailer (1954) summarized the literature, both bionomic and economic, pertaining to this species. 16 specimens examined. Collection dates from 16 May to 15 September. Distribution: Clinton, Kalamazoo, Kent, and Washtenaw counties.

Subfamily SEHIRINAE Amyot & Serville, 1843 Genus SEHIRUS Amyot & Serville, 1843 Sehirus cinctus albonotatus Dallas, 1851. – Froeschner (1960) reported this species from Michigan. Stoner’s (1922) record of Sehirus cinctus remains unplaced to subspecies, but the author has examined actual individuals of S. c. albonotatus from Cheboygan County (see below). This subspecies also has been collected from Mackinac Island (Mackinac County). This cydnid, along with the nominate subspecies below, are commonly referred to as the white- margined burrower bug. 71 specimens examined. Collection dates from 16 May to 15 September. Distribution: Alcona, Charlevoix, Cheboygan, Chippewa, Delta, Dickinson, Emmet, Gogebic, Iosco, Iron, Kalamazoo, Luce, Mackinac, Manistee, Marquette, Menominee, Midland, Montmorency, Muskegon, Newaygo, Oceana, Ontonagon, and Tuscola counties. Sehirus cinctus cinctus (Palisot de Beauvois), 1811. – Stoner (1922) reported Sehirus cinctus swept from weeds in a rye field in Cheboygan County, although it is not known to what subspecies his record applies. He did not indicate the presence of subapical corial spots; thus, one might assume the record refers to the nominate subspecies. Nevertheless, the record was not included for lack of certainty. This subspecies has been taken on willow (Salix sp.) in Ber- rien County and black raspberry (Rubus occidentalis L.) in Livingston County. It also has been collected from a Malaise trap in Livingston County, a rotary trap in a wheat field in Berrien County, and a rotary trap in an oat field in both Berrien and Kalamazoo counties. McDonald (1968a) and Sites and McPherson (1982) provided notes on the life history of this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. 93 specimens examined. Collection dates from 13 April to 13 September. Distribution: Allegan, Berrien, Cass, Eaton, Grand Traverse, Gratiot, Ingham, Jackson, Kalamazoo, Kent, Livingston, Mackinac, Mason, Monroe, Oakland, Osceola, Saint Joseph, Shiawassee, Washtenaw, and Wayne counties. 278 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Family ACANTHOSOMATIDAE Signoret, 1863 The acanthosomatids superficially resemble and might easily be mistaken for pentatomids, and in the past, the family had been afforded subfamilial or tribal rank within the Pentatomidae. The group, however, has been shown to occupy a more basal position in the pentatomoid phylogeny (Grazia et al. 2008). The species, perhaps unlike other pentatomoids, seem to prefer shrubs and trees over forbs, and Schaefer and Ahmad (1987) discussed some of the host plants of this family. These insects also are more abundant in the northern United States and Canada (Froeschner 1988a) than in the southern states. Members of this group are notable for reports of extended maternal care (Frost and Haber 1944). Acanthosomatidae is a small family, contrasting the great diversity of some of the other pentatomoid groups. Kumar (1974) revised the Acanthosomatidae on a global scale and supplied a generic key, whereas Rolston and Kumar (1974) provided keys to the species of the Western Hemisphere. Thomas (1991) reviewed the North American species of the family. Froeschner’s (1988a) catalog is the most current for the United States fauna, and 5 species in 2 Holarctic genera occur in the United States, all belonging to the nominate subfamily. Both genera are represented in Michigan, and 3 of the 5 species are known from the state; the northwestern Elasmostethus interstinctus (Linnaeus), 1758 and southwestern Elasmucha cordillera Thomas, 1991 are absent. The key provided is modified from McPherson’s (1970) treatment.

Key to the Acanthosomatidae of Michigan 1 Ostiolar canal short, reaching only middle of metapleuron; inner angles of posterior margin of pronotum produced posteriorly...... Elasmucha lateralis 1’ Ostiolar canal long, much surpassing middle of metapleuron; inner angles of posterior margin of pronotum not produced posteriorly (Elas- mostethus)...... 2 2 (1’) Antennae piceous or shining black with the incisures pale; pronotal punctures fine, concolorous anteriorly, dark narrow series posteriorly; humeri to posterior angles blackish...... Elasmostethus atricornis 2’ Antennae pale except apical segment more or less darker; pronotal punctures coarse, dark, almost foveolate, widely separated; humeri to posterior angles more or less pale or reddish...... Elasmostethus cruciatus

Subfamily ACANTHOSOMATINAE Signoret, 1863 Genus ELASMOSTETHUS Fieber, 1861 Elasmostethus atricornis (Van Duzee), 1904. – This species was recorded from Michigan by McPherson (1970). Jones and McPherson (1980) and Carter and Hoebeke (2003) provided bionomic information for this acanthosomatid. 30 specimens examined. Collection dates from 10 July to 15 August. Distribution: Alcona, Cheboygan, Clare, Gogebic, Grand Traverse, Iosco, Lapeer, Manistee, Midland, and Muskegon counties. Elasmostethus cruciatus (Say), 1831. – Occasionaly referred to as the red- cross shield bug, E. cruciatus was recorded from Michigan by Stoner (1922), who collected one female sweeping vegetation along a low swampy lakeshore. It also has been collected on Drummond Island (Chippewa County), Isle Royale (Keweenaw County), and Mackinac Island (Mackinac County). Jones and McPherson (1980) provided life history data on this species in South Carolina. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. 67 specimens examined. Collection dates from 8 May to 7 October. 2012 THE GREAT LAKES ENTOMOLOGIST 279 Distribution: Cheboygan, Chippewa, Delta, Dickinson, Emmet, Houghton, Huron, Kalamazoo, Keweenaw, Lapeer, Leelanau, Mackinac, Manistee, Mar- quette, Midland, Ogemaw, Ontonagon, and Roscommon counties.

Genus ELASMUCHA Stål, 1864 Elasmucha lateralis (Say), 1831. – This species was recorded from Michi- gan by Stoner (1922) as Meadorus lateralis; McPherson (1970) also treated it under that binomial. Stoner (1922) took several individuals, all in wet or swampy situations, including one on a lily pad. Label data indicate that it also has been taken on flowers of meadow sweet (Spiraea alba Du Roi) in Montcalm County. However, E. lateralis has a well-established association with birch and beech (Jones and McPherson 1980), and these records seem to represent incidental encounters. Additional label data indicate that twenty- three individuals were taken in beach drift in early June in Baraga County. This acanthosomatid also has been collected on Isle Royale (Keweenaw County). Jones and McPherson (1980) provided life history information on this species in South Carolina. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. 180 specimens examined. Collection dates from 4 May to 5 October. Distribution: Alcona, Alger, Antrim, Baraga, Charlevoix, Cheboygan, Chippewa, Clare, Delta, Emmet, Gladwin, Gogebic, Houghton, Ingham, Iosco, Iron, Isabella, Kalamazoo, Keweenaw, Leelanau, Luce, Mackinac, Manistee, Marquette, Mason, Missaukee, Montcalm, Oceana, Ontonagon, Roscommon, and Saint Joseph counties.

Family PENTATOMIDAE Leach, 1815 The Pentatomidae are one of the largest and most diverse of heteropteran families. Comprising approximately 224 species in 64 genera in the United States (Froeschner 1988c), the stink bugs, although varied in morphology, still generally retain the recognizable stink bug habitus. Many species, whether primarily herbivorous or predaceous, are generalist feeders. Adults usually overwinter, often becoming conspicuous guests in homes. Eggs generally are laid in the spring; most are uni- or bivoltine in Michigan, although multivolt- inism is known to occur in a few species (McPherson 1982). Many pentatomids frequent lights at night, occasionally in large aggregations. The Asopinae, or predatory stink bugs, are members of this family and are easily recognized by the free first segment of the rostrum. Also included are the Podopinae (previously the Graphosomatinae), or turtle bugs, a group of generally paludicolous pentatomids with a greatly enlarged scutellum, similar to the Scutelleridae. The economic significance varies from species to species, with many being innocuous and others encompassing some of the worst pests of commercial crops known to mankind. McPherson and McPherson (2000) provided a wealth of information on the economic importance of the family in North America, and De Clercq (2000) and Panizzi et al. (2000) discussed the positive impact of the asopines and the negative impact of the herbivorous pentatomids, respectively, on a world scale. Froeschner (1988c) provided the most current catalog to the species found north of Mexico. Five subfamilies are represented in the United States, with the majority of the species belonging to the nominate Pentatominae. Two small subfamilies, the Discocephalinae and Edessinae, are represented each by two species in the United States; they do not occur in Michigan. Esselbaugh (1948) provided bionomic information for many Midwestern species. Thomas (1992) treated the Asopinae of the New World. Barber and Sailer (1953) and Davidová-Vilímová and McPherson (1994) dealt with the classification of the 280 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Podopinae. The Pentatominae were treated in a series of papers by Rolston and McDonald (1979, 1980, 1984), and Rolston et al. (1980). In Michigan, 49 species in 29 genera are known. The following key is synthesized from McPherson (1970) and the other taxonomic works mentioned above. Although it would be more desirable to shape the key to better follow the phylogeny of the group, I have not been able to find a more satisfactory treatment of the tribal classification than is offered by Rolston and McDonald (1979). Additionally, three species (Perillus strigipes (Asopinae), Brochymena carolinensis (Pentatominae: Halyini), and Murgantia histrionica (Pentatominae: Pentatomini)) reported from Michigan by Furth (1974) but based on unconfirmed records, are included in the key to facilitate identification of legitimate records.

Key to the Pentatomidae of Michigan 1 First rostral segment directed away from head, incrassate, free, with only base between bucculae which converge ventrally (Asopinae)...... 2 1’ First rostral segment not directed away from head, slender, lying between bucculae which are subparallel...... 15 2 (1) Scutellum greatly enlarged, U-shaped, covering most of hemelytra and abdomen; frena about one-fourth to one-third length of scutellum; protibia greatly dilated...... Striretrus anchorago 2’ Scutellum not greatly enlarged, more or less triangular, leaving hemelytra and abdomen broadly exposed; frena about one-half or more length of scutellum; protibia not distinctly dilated...... 3 3 (2’) Apical part of scutellum (tongue) enlarged, about as wide as corium...... 4 3’ Apical part of scutellum small, much narrower than corium...... 8 4 (3) Juga distinctly longer than tylus, convergent and contiguous in front; abdominal venter without anteriorly projecting process; profemur without subdistal spine...... Rhacognathus americanus 4’ Juga subequal to tylus; abdominal venter with prominent stout anterior directed tubercle; profemur with small to large subdistal spine (Perillus)...... 5 5 (4’) Profemur with subdistal spine obsolescent, spine not longer than wide...... 6 5’ Profemur with distinct subdistal spine which is longer than wide.....7 6 (5) Pronotum orange or yellow with uninterrupted medial transverse black stripe or pronotum mostly black except for margins, never with midlongitudinal stripe; scutellum with U-shaped submarginal band of yellow or with apex pale, never chevron-shaped...... Perillus exaptus 6’ Pronotum black with yellow or orange midlongitudinal stripe and/or with yellow to orange chevron on scutellum ...... Perillus strigipes 7 (5’) Pronotum usually with pale transverse fascia in middle; abdominal venter with submarginal row of black spots which enclose spiracles...... Perillus bioculatus 7’ Pronotum usually without pale transverse fascia; abdomen lacking black submarginal ventral spots, spiracles enclosed within yellow area...... Perillus circumcinctus 8 (3’) Second abdominal sternite unarmed; ostiole of scent gland without elevated ruga, evaporatorium reduced; dorsal color metallic blue; size small, length usually less than 8 mm...... Zicrona caerulea 2012 THE GREAT LAKES ENTOMOLOGIST 281 8’ Second abdominal sternite produced cephalad as spine or tubercle; ostiole of scent gland attended by elevated ruga and surrounded by distinct evaporatorium; dorsal color brown, reddish, or grey, not metallic blue; size larger, length greater than 8 mm ...... 9 9 (8’) Juga usually slightly longer than tylus; males with glandular patches of silky hairs on abdominal venter; female basal gonocoxites separated by intervening plate or contiguous, if contiguous then inner margins of juga distinctly convergent; size larger, length usually 14 mm or more (Apoecilus)...... 10 9’ Juga equal to tylus; males without glandular patches of silky hairs on abdominal venter; female basal gonocoxites contiguous; inner margins of juga parallel; size smaller, length usually 12 mm or less (Podisus)..... 11 10 (9) Anterior half of pronotum usually with six dark spots, two submedian spots between pairs of submarginal spots; basal gonocoxites of female convergent apically, mesal plate between them triangular; upper arm of male paramere curvilinear, not bent, about equal in diameter to lower arm...... Apoecilus bracteatus 10’ Anterior half of pronotum usually with only four dark spots, two submarginal spots on each side; basal gonocoxites of female not convergent apically, mesal plate between them quadrangular; upper arm of male paramere bent or more slender in diameter than lower arm...... Apoecilus cynicus 11 (9’) Anterolateral margin of pronotum straight; hemelytral membrane clear (sometimes with faint stripe); [humeri rounded]...... Podisus placidus 11’ Anterolateral pronotal margin concave; hemelytral membrane with dark blotch or stripe...... 12 12 (11’) Humeral angles acute to spinose; femora with pair of anteapical spots; seventh abdominal sternite usually with median spot...... Podisus maculiventris 12’ Humeral angles rounded or obtusely angular; femora variably maculate or immaculate; abdominal sternites immaculate or with row of spots...... 13 13 (12’) Basal ventral abdominal spine long, protruding between and attaining anterior margin of metacoxae...... Podisus neglectus 13’ Basal ventral abdominal spine short, not attaining anterior margin of metacoxae...... 14 14 (13’) Metafemora immaculate; abdominal venter without median row of spots...... Podisus brevispinus 14’ Metafemora densely spotted; abdominal venter usually with median row of spots ...... Podisus serieventris 15 (1’) Scutellum greatly enlarged, U-shaped, covering most of hemelytra and abdomen; frena much less than one-fourth as long as scutellum (Podopinae: Amaurochrous)...... 16 15’ Scutellum not greatly enlarged, more or less triangular, leaving hemelytra and abdomen broadly exposed; frena at least one-fourth as long as scutellum (Pentatominae)...... 17 16 (15) Juga distinctly longer than tylus, often contiguous in front; fifth antennal segment subequal to second, third and fourth combined; scutellum more than twice as long as head and pronotum combined...... Amaurochrous brevitylus 282 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

16’ Juga subequal to tylus; fifth antennal segment slightly longer than third and fourth combined; scutellum one-fourth longer than head and pronotum combined...... Amaurochrous cinctipes 17 (15’) Juga with lateral tooth near apex; hemelytral membrane with arbo- rescent dark markings (Hyalini)...... 18 17’ Juga without lateral tooth near apex; hemelytral membrane immaculate or with streaks or spots...... 20 18 (17) Humeral angles subquadrate with large teeth...... Parabrochymena arborea 18’ Humeral angles subtriangular with small teeth (Brochymena)...... 19 19 (18’) Juga subequal to tylus or exceeding tylus by less than jugal width at apex of tylus...... Brochymena carolinensis 19’ Juga distinctly longer than tylus, often convergent and contiguous in front...... Brochymena quadripustulata 20 (17’) Body broad, flat with margins explanate, broadest behind middle; [second antennal segment at least 1.5 times length of third]; head nearly as wide as base of scutellum (Sciocorini)...... Sciocoris microphthalmus 20’ Body usually broadest before middle, margins not flattened and explanate (except Menecles, in this case second antennal segment less than 1.5 times length of third); head usually narrower than base of scutellum or, if nearly as wide, not flat above (Pentatomini)...... 21 21 (20’) Second abdominal sternite produced cephalad as spine or tubercle...... 22 21’ Second abdominal sternite unarmed...... 27 22 (21) Juga surpassing tylus, convergent and contiguous in front...... Dendrocoris humeralis, females (in part) 22’ Juga subequal to tylus...... 23 23 (22’) Abdominal process spinous, reaching between metacoxae; metasternum unmodified, not in apposition with abdominal spine Chinavia( )...... 24 23’ Abdominal process tuberculate, not extending between metacoxa; metasternum subelevated, notched posteriorly, in apposition with abdominal tubercle (Banasa)...... 25 24 (23) Anterolateral margins of pronotum straight or nearly so; width of head across eyes equal or subequal to length; rostrum reaching between metacoxae; connexival spots at least on basal segments expanding onto laterotergite...... Chinavia hilaris 24’ Anterolateral margins of pronotum broadly rounded; head wider across eyes than long; rostrum reaching only to mesocoxae; connexiva immaculate or black mark at posterolateral angles of segments confined to edge of sternite...... Chinavia pensylvanica 25 (23’) Pronotum, disc of scutellum, and corium concolorous, usually grey or sordid brown; rostrum long, surpassing posterior margin of tubercle- bearing abdominal sternite...... Banasa sordida 25’ Pronotum bicolored, anterior half lighter than posterior half and/or corium with pale or reddish coloration contrasting with greenish scutellum; rostrum short, not reaching posterior margin of tubercle-bearing abdominal sternite...... 26 26 (25’) Metapleuron without or with small inconspicuous spot; juga without pigmented punctations...... Banasa calva 26’ Metapleuron with distinct stigmatose spot; juga usually with pigmented punctations...... Banasa dimidiata 2012 THE GREAT LAKES ENTOMOLOGIST 283 27 (21’) Margins of propleura before coxae produced, anterior margin covering base of head from eyes mesad nearly to rostrum; [metathoracic ostiole normally with distinct auricle, existing, at most, as short abruptly ending canal, rounded on inner margin; scutellum equal to or longer than corium with apical third wider than apex of corium and with tip broadly rounded]...... 28 27’ Propleura not produced...... 30 28 (27) Base of antennae hidden from lateral view by anterior production of propleuron; pronotum longitudinally tricarinate...... Aelia americana 28’ Antenniferous tubercles not covered by anterior production of propleuron; pronotum only midlongitudinally carinate (Neottiglossa)...... 29 29 (28’) Head entirely black, with deep dense punctures; vertex without pale line extending caudad onto pronotum...... Neottiglossa trilineata 29’ Head not black, without deep dense punctures; vertex with median pale line extending caudad onto pronotum...... Neottiglossa undata 30 (27’) Metathoracic ostiole unattended by sulcus or ruga; brightly colored black, red, and yellow species...... Murgantia histrionica 30’ Sulcus or ruga of various lengths extending laterad from metathoracic ostiole; species not brightly or contrastingly colored...... 31 31 (30’) Metathoracic ostiole without auricle, existing as tapering canal, V- shaped on inner margin...... 32 31’ Metathoracic ostiole normally with distinct auricle, existing, at most, as short abruptly ending canal, rounded on inner margin...... 41 32 (31) Canal of metathoracic ostiole not extending beyond middle of metapleuron...... 33 32’ Canal of metathoracic ostiole extending beyond middle of metapleuron ...... 35 33 (32) Head and pronotum glabrous; green species...... Chlorochroa persimilis 33’ Head and usually pronotum hirsute; tan or brownish species (Tricho- pepla)...... 34 34 (33’) Head broadly rounded or truncate apically, ratio of width to length 1:1 or less, dorsal surface with few hairs; pseudoclaspers with few lateral spines at most...... Trichopepla atricornis 34’ Head distinctly elongate, tapering apically, ratio of width between eyes to length (apex to line across base of eyes) 1:1.3, dorsal surface often densely pilose; pseudoclaspers with numerous stout spines on surface...... Trichopepla semivittata 35 (32’) Size large, length greater than 12 mm; third and fourth antennal segments distinctly banded; dorsal and ventral surface of pronotum with metallic green punctures...... Halyomorpha halys 35’ Size smaller, length 10 mm or less; third and fourth antennal segments not distinctly banded; dorsal and ventral surface of pronotum without metallic green punctures ...... 36 36 (35’) Juga shorter or subequal to tylus (Thyanta)...... 37 36’ Juga longer than tylus...... 38 37 (36) Anterolateral margins of pronotum and apical angles of abdominal segments pale or concolorous; pronotal cicatrices without piceous spots on inner angles; elytra usually provided with scattered pale spots...... Thyanta custator accerra 284 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

37’ Anterolateral margin of pronotum and apical angles of abdominal segments with black spots; pronotal cicatrices with piceous spot in inner angles; elytra usually without scattered pale spots...... Thyanta calceata 38 (36’) Anterolateral margins of pronotum strongly concave...... Dendrocoris humeralis, males (in part) 38’ Anterolateral margins of pronotum, at most, weakly concave (Holcoste- thus)...... 39 39 (38’) Abdominal venter dark chocolate brown, margins occasionally yellow...... Holcostethus mcdonaldi 39’ Abdominal venter yellow or reddish brown, with or without black markings...... 40 40 (39’) Reddish brown species with distinctive black zig-zag markings on abdominal venter; juga not contiguous in front of tylus...... Holcostethus fulvipes 40’ Brown colored species without distinct markings on abdominal venter; juga contiguous in front of tylus...... Holcostethus limbolarius 41 (31’) Body hirsute; posterior margins of humeral angles emarginate...... Prionosoma podopioides 41’ Body glabrous; humeral angles prominent or rounded but not emarginate...... 42 42 (41’) Scutellum equal to or longer than corium, apical third wider than apex of corium, apex broadly rounded...... 43 42’ Scutellum shorter than corium, apical third narrower than apex of corium, apex not broadly rounded...... 44 43 (42) Head horizontal or slightly declivent; tylus distinctly elevated above juga; dull brownish species...... Coenus delius 43’ Head more deflexed; tylus scarcely elevated above juga; black and red species...... Cosmopepla lintneriana 44 (42’) Dorsal surface of tibia rounded for most of its length...... 45 44’ Dorsal surface of tibia sulcate for most of its length...... 46 45 (44) Humeral angles rounded; black species with distinct white scutellar markings...... Mormidea lugens 45’ Humeral angles distinctly spined; tan or brownish species...... Oebalus pugnax pugnax 46 (44’) Anterolateral pronotal margins entire...... 47 46’ Anterolateral margins of pronotum crenulate or denticulate (Euschis- tus)...... 49 47 (46) Head deeply inserted into pronotum, imaginary line drawn across anterior limits of pronotum bisecting eyes near middle; margins of pronotum strongly explanate; rostrum reaching second abdominal sternite...... Menecles insertus 47’ Head not deeply inserted into pronotum, imaginary line drawn across anterior limits of pronotum passing near posterior margin of eyes; margins of pronotum not explanate; rostrum not surpassing metacoxae...... 48 48 (47’) Hemelytral membrane with numerous small dark spots; veins nearly obscure, few, simple or bifurcate; posterior margin of bucculae sloping abruptly in lateral view; anterolateral margins of pronotum straight to slightly concave; smaller, length 6-8.5 mm...... Mcphersonarcys aequalis 2012 THE GREAT LAKES ENTOMOLOGIST 285 48’ Hemelytral membrane without spots, unmarked or brown except along veins; venation reticulate; posterior margin of bucculate roundly truncate in lateral view; anterolateral margins of pronotum broadly rounded; larger, length 9-11.5 mm...... Hymenarcys nervosa 49 (46’) Pronotum with irregular raised impunctate transverse line between humeral angles...... Euschistus ictericus 49’ Pronotum without raised impunctate line between humeral angles.....50 50 (49’) Incisures of abdominal sternites without black spots near anterolateral angle...... Euschistus variolarius 50’ Incisures of abdominal sternites with black spot near anterolateral angle...... 51 51 (50’) Abdominal sternites with one or more medial black spots (Euschistus tristigmus)...... 52 51’ Abdominal sternites without black medial spots...... 53 52 (51) Humeral angles rounded; apical half of fourth antennal segment and nearly all of fifth fuscous...... Euschistus tristigmus luridus 52’ Humeral angles acute or spinose; antennae entirely rufous or testa- ceous...... Euschistus tristigmus tristigmus 53 (51’) Anterolateral margins of pronotum straight or nearly so, pale margin bordered within by well-defined line of black punctures...... Euschistus politus 53’ Anterolateral margins of pronotum distinctly sinuate, pale margin not bordered within by well-defined line of black punctures (Euschistus servus)...... 54 54 (53’) Juga clearly exceeding apex of tylus...... Euschistus servus euschistoides 54’ Juga equaling or slightly exceeding apex of tylus...... Euschistus servus servus

Subfamily ASOPINAE Spinola, 1850 Genus APOECILUS Stål, 1870 Apoecilus bracteatus (Fitch), 1856. – This species was reported from Michigan by Hart (1919) as Apateticus crocatus (Uhler), 1897; Thomas’s (1992) treatment of that name as a junior synonym of A. bracteatus is followed. Stoner (1922) and Hussey (1922a) each reported Apateticus bracteatus from Michigan. Stoner (1922) found this species to be more common than the the following species and swept several individuals from willow (Salix sp.) and collected some from beach drift. Hussey found A. bracteatus to be much less common than its Michigan congener, but also took several individuals from beach drift. In Michigan, the members of this genus are the largest predatory stink bugs as well as among the largest pentatomoids. Evans and Root (1980) reported biological information for this species. De Clerq (2000) discussed this as one of the less economically important species. McDonald (1966) described the genitalia. Thomas (1992) raised Apoecilus to full generic rank. 53 specimens examined. Collection dates from 6 June to 3 October. Distribution: Alger, Allegan, Barry, Berrien, Charlevoix, Cheboygan, Chippewa, Delta, Emmet, Gladwin, Gratiot, Houghton, Huron, Iosco, Isabella, Keweenaw, Livingston, Luce, Mackinac, Marquette, Midland, Oakland, Oge- maw, Schoolcraft, Washtenaw, and Wayne counties. Apoecilus cynicus (Say), 1832. – This species was reported from Michigan by Townsend (1890) (as Podisus cynicus). Stoner (1922) collected this species from beach drift. As Apateticus cynicus, Hussey (1922a) reported: 286 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

“Nymphs which probably should be referred to this species were common on the maples...and one adult female was beaten from a maple there...this form was by far the most abundant of the beach-drift of early July; it was estimated that 80 per cent of the Hemiptera washed up between July 7 and July 15 were A. cynicus, and on some days the number of individuals of this species on the beach was estimated at about 250 per mile.” It has also been taken beating the edge of wooded areas in Saint Joseph County. In Michigan, the members of this genus are the largest predatory stink bugs as well as among the largest pentatomoids. Jones and Coppel (1968) reported biological information for this species. De Clerq (2000) discussed this as one of the less economically important species. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. Thomas (1992) raised Apoecilus to full generic rank. 89 specimens examined. Collection dates from 18 June to 12 November. Distribution: Alger, Allegan, Berrien, Cheboygan, Crawford, Gladwin, Houghton, Huron, Ingham, Iosco, Isabella, Keweenaw, Lenawee, Livingston, Midland, Muskegon, Oakland, Oceana, Oscoda, Saint Joseph, Shiawassee, Van Buren, and Washtenaw counties.

Genus PERILLUS Stål, 1862 Perillus bioculatus (Fabricius), 1803. – The two-spotted stink bug was reported from Michigan by Yothers (1911) (as Perillus claudus). Hussey (1922a) collected this species from the beach-drift. Stoner (1922) swept an individual from a low grassy area in a rye field and also noted a dead, presumably overwintering individual taken indoors. It was taken in a house in Livingston County on 12 March as well as in a museum on 18 October and in a house on 12 December in Washtenaw County, suggesting it may come into contact with humans as a frequent overwinterer, which also support Stoner’s (1922) observation. McDaniel (1924) also discussed the economic impact of this species in Michigan, like Yothers (1911), as specifically related to the Colorado potato beetle (Leptinotarsa decimlineata (Say), 1824); Froeschner (1988c) cast doubt on this instance of host specificity in the predaceous pentatomid. Knight (1922, 1924) discussed the life history and variables affecting the color and color pattern of this species. De Clerq (2000) discussed the economic impact of this species. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. Knight (1952) reviewed and keyed the genus. 91 specimens examined. Collection dates from 11 March to 25 December. Distribution: Alger, Antrim, Arenac, Bay, Benzie, Berrien, Branch, Cass, Cheboygan, Clinton, Eaton, Emmet, Huron, Ingham, Kalamazoo, Kent, Lapeer, Livingston, Mackinac, Macomb, Marquette, Menominee, Midland, Missaukee, Montcalm, Muskegon, Oakland, Oceana, Ottawa, Presque Isle, Saint Clair, Saint Joseph, Van Buren, Washtenaw, Wayne, and Wexford counties. Perillus circumcinctus Stål, 1862. – McPherson (1970) reported this species from Michigan. Evans (1982a, 1982b, 1982c) provided biological information. Knight (1952) reviewed and keyed the genus. 13 specimens examined. Collec- tion dates from 10 June to 31 July. Distribution: Cheboygan, Houghton, Mecosta, Oakland, Roscommon, Van Buren, Washtenaw, and Wayne counties. Perillus exaptus (Say), 1825. – This species was reported from Michigan by Stoner (1922). It has been taken in a window pane trap in Montmorency County. Knight (1952) reviewed and keyed the genus. 24 specimens examined. Collection dates from 17 April to 7 August. 2012 THE GREAT LAKES ENTOMOLOGIST 287 Distribution: Charlevoix, Cheboygan, Chippewa, Clare, Crawford, Dickin- son, Delta, Emmet, Iosco, Lake, Livingston, Mackinac, Marquette, Montmorency, Newaygo, Osceola, Roscommon, and Wexford counties. Perillus strigipes (Herrich-Schaeffer), 1853. – Furth (1974) listed this species among the Michigan fauna based on specimens purportedly housed in the UMMZ, but McPherson (1970, 1979b) could not locate the specimens. They remain unfound. The species, however, is included in the key to facilitate identification of legitimate records. Nevertheless, it should be dropped from the Michigan faunal list until the specimens are found or additional material is collected. This species was excluded from Knight’s review (1952) of Perillus as the P. strigipes was included in the genus Mineus Stål, 1867, until Thomas (1990) synonymized the names.

Genus PODISUS Herrich-Schaeffer, 1851 Podisus brevispinus Phillips, 1982. – Stoner (1922) and Hussey (1922a) reported this species from Michigan (both as Podisus modestus). Hussey (1922a) took the species “in the same situations about the Warren Woods as was [P. maculiventris], but not found either in the dune region or in the beach drift.” Stoner (1922), however, did take this species in beach drift, and additional label data indicate it has been on beaches as well as from unspecified flowers in Cheboygan County. This species has been taken on flowers of meadow sweet (Spiraea alba Du Roi) in Montcalm County. It also has been collected on Drummond Island (Chippewa County). This species previously has been recorded in the United States as Podisus modestus (Dallas), 1851, but this name was shown to refer to P. maculiventris. Phillips (1983) clarified the status of Arma modestus Dallas, 1851 and renamed this taxon, which was accepted by Thomas (1992). Tostowaryk (1971) described the life history of this species in Quebec. Evans (1985) provided a key for separating the nymphs of this species from several of its congeners. De Clerq (2000) discussed this as one of the less economically important species. 175 specimens examined. Collection dates from 8 April to 23 October. Distribution: Alger, Allegan, Baraga, Barry, Bay, Benzie, Berrien, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gogebic, Gratiot, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Livingston, Luce, Mackinac, Marquette, Midland, Missaukee, Monroe, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Roscommon, Saginaw, Schoolcraft, Shiawassee, Washtenaw, and Wayne counties. Podisus maculiventris (Say), 1832. – Commonly referred to as the spined soldier bug, P. maculiventris was reported from Michigan by Townsend (1890) based on “larvae, probably [Podisus spinosus], often found with imagos on various plants in early summer.” Stoner (1922) suggested this species might be found in low, moist, grassy areas and took several specimens from willow (Salix sp.) and in beach drift. Hussey (1922a) reported this species “[r]ather common on various bushes and in the grass about Warren Woods...and also found in the dune region. This was one of the most common species in the beach drift.” This species is the most common predatory pentatomid in Michigan and has been taken in a variety of circumstances. Label data describe one collection of P. maculiventris as “Sweeping fallow roadside. Solidago dominant. Some shrubs.” in Isabella and Jackson counties. In Monroe County, it was taken under a board on a beach, and it has been taken in open woods in Washtenaw County. This species has been taken from a rotary trap in a wheat field in Berrien County as well as white and ultraviolet lights in Clinton and Livingston counties. It has 288 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 been taken from cultivated strawberries (Fragaria sp.) in Berrien County. A specimen from Kalamazoo County has been pinned with its moth prey, and it has been taken “feeding on Malacosoma sp. [Lepidoptera: Lasiocampidae] larvae” in Livingston County. Mukerji and LeRoux (1969) and McPherson (1980b) supplied information on the feedings habits of this species. Evans (1985) provided a key for separating the nymphs of this species from several of its congeners. De Clerq (2000) discussed the economic impact of this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. 426 specimens examined. Collection dates from 12 April to 13 November. Distribution: Allegan, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Clare, Clinton, Crawford, Delta, Eaton, Genesee, Gladwin, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Mid- land, Missaukee, Monroe, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Ottawa, Roscommon, Saginaw, Saint Joseph, Schoolcraft, Shiawassee, Tuscola, Van Buren, Washtenaw, and Wayne counties. Podisus neglectus (Westwood), 1837. – Blatchley (1926) recorded this species from Michigan (as Podisus fretus Olsen, 1916), which McPherson (1970, 1979b) tentatively included and later confirmed. Thomas (1992) synonymized P. fretus under Westwood’s taxon. 1 specimen examined. Collection dates from 10 June to 10 September. Distribution: Houghton and Ottawa counties. Podisus placidus Uhler, 1870. – This species was reported from Michigan by Townsend (1890) taken on “flowers of golden rod.” Stoner (1922) found this to be the most common Podisus species around Douglas Lake, taking it from swampy regions, burn-over areas, and beach drift as well as from balsam poplar (Populus balsamifera L.), white cedar (Thuja occidentalis L.), and willow (Salix sp.). It has been collected in a Malaise trap at 6 feet height in Manistee County and a window pane trap in Oscoda County. This asopine has been taken from eastern black oak (Quercus velutina Lamb.) in Livingston County. Oetting and Yonke (1971) gave bionomic information. Evans (1985) provided a key for separating the nymphs of this species from several of its congeners. De Clerq (2000) discussed this as one of the less economically important species. 92 specimens examined. Collection dates from 26 April to 9 October. Distribution: Alcona, Allegan, Alpena, Barry, Cass, Charlevoix, Che- boygan, Clare, Clinton, Crawford, Grand Traverse, Gratiot, Huron, Ingham, Iron, Kalamazoo, Kalkaska, Lake, Lapeer, Livingston, Mackinac, Manistee, Marquette, Midland, Montcalm, Montmorency, Newaygo, Oakland, Oscoda, Otsego, Roscommon, Saint Joseph, Tuscola, Van Buren, Washtenaw, Wayne, and Wexford counties. Podisus serieventris Uhler, 1871. – Hart (1919) reported this species from Michigan (as Apateticus (Podisus) serieventris). This species has been taken on tamarack (Larix laricina (Du Roi) Koch) and red pine (Pinus resinosa Sol. ex. Aiton) in Livingston County; P. serieventris has also been taken on tamarack in Oakland County. Prebble (1933) reported biological information. Evans (1985) provided a key for separating the nymphs of this species from several of its congeners. De Clerq (2000) discussed this as one of the less economically important species. 73 specimens examined. Collection dates from 12 April to 8 October. Distribution: Alcona, Cheboygan, Chippewa, Crawford, Emmet, Houghton, Huron, Ingham, Iosco, Kalamazoo, Keweenaw, Lake, Leelanau, Livingston, Mackinac, Oakland, Oceana, Saint Joseph, and Washtenaw counties. 2012 THE GREAT LAKES ENTOMOLOGIST 289 Genus RHACOGNATHUS Fieber, 1861 Rhacognathus americanus Stål, 1870. – This species was reported from Michigan by Hussey (1922a) who noted “[o]ne mutilated individual of this very rare species was found in old beach-drift.” Steyskal (1938) provided additional records. The long convergent juga will easily separate this species from all other Michigan asopines. McDonald (1966) described the genitalia. 4 specimens examined. Collection dates from 25 May to 18 July. Distribution: Berrien, Clare, Livingston, and Midland counties.

Genus STIRETRUS Laporte, 1833 Stiretrus anchorago (Fabricius), 1775. – Commonly referred to as the anchor stink bug, this species was reported from Michigan by McPherson (1970). Much variation exists in the color and color pattern of this species, and many color forms were subsequently described either as separate species or as varieties of S. anchorago. I follow Thomas (1992) in essentially ignoring all the color forms; nevertheless, only individuals that would fall under Stiretrus anchorago var. fimbriatus(Say), 1828 occur in Michigan. The enlarged scutellum is unique among the Michigan pentatomids, differing from the podopines, and will easily separate it from all other taxa. This species has been taken from wrinkleleaf goldenrod (Solidago rugosa Mill.) and early goldenrod (S. juncea Aiton) in Cheboygan County. Waddill and Shepard (1974) described the biology. De Clerq (2000) discussed this as one of the less economically important species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. 48 specimens examined. Collection dates from 18 May to 11 September. Distribution: Arenac, Barry, Berrien, Cheboygan, Clare, Clinton, Delta, Dickinson, Gladwin, Grand Traverse, Gratiot, Huron, Ingham, Iosco, Isa- bella, Jackson, Kalkaska, Kent, Menominee, Midland, Missaukee, Montcalm, Montmorency, Newaygo, Ogemaw, Oscoda, Otsego, Roscommon, Tuscola, and Wexford counties.

Genus ZICRONA Amyot & Serville, 1843 Zicrona caerula (Linnaeus), 1758. – Hart (1919) reported this species from Michigan. The small habitus and shiny deep blueish color will render this species instantly recognizable within the Michigan pentatomid fauna. De Clerq (2000) discussed this as one of the less economically important species. McDonald (1966) described the genitalia. 2 specimens examined. Collection dates from 29 May to 18 June. Distribution: Houghton and Schoolcraft counties.

Subfamily PENTATOMINAE Leach, 1815 Tribe HALYINI Amyot & Serville, 1843 Genus BROCHYMENA Amyot & Serville, 1843 Brochymena carolinensis (Westwood), 1837. – Furth (1974) listed this species among the Michigan fauna based on specimens purportedly housed in the UMMZ, but McPherson (1970, 1979b) could not locate the specimens. They remain unfound. The species, however, is included in the key to facilitate identification of legitimate records. Nevertheless, it should be dropped from the Michigan faunal list until the specimens are found or additional material is collected. 290 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Brochymena quadripustulata (Fabricius), 1775. – The author assumes that Townsend (1890) reported this species from Michigan as Brochymena annulata since only two species of the genus are known from Michigan (and B. quadripustulata quite commonly so); McPherson (1980a) also had come to this conclusion. Nevertheless, B. quadripustulata, commonly referred to as the four-humped stink bug, was unequivocally reported by McPherson (1970) in his treatment of the Michigan pentatomoids. Label data describe one collection in Isabella County as “Sweeping fallow roadside. Solidago dominant. Some shrubs.” It also has been taken on tamarack (Larix laricina Du Roi) in Oakland County and wild grape (Vitis riparia Michx.) in Livingston County. Ruckes (1946) reported this species as occasionally predaceous on soft-bodied larvae. Cuda and McPherson (1976) provided notes on the biology and laboratory rearing for this species. Gamboa and Alcock (1973) described the mating behavior, and McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Ruckes (1946) keyed the genus, and Larivière (1992) keyed those species excluded from the arborea group. 216 specimens examined. Collection dates from 20 March to 30 November. Distribution: Allegan, Barry, Bay, Berrien, Cass, Charlevoix, Chippewa, Clare, Clinton, Crawford, Eaton, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Ingham, Ionia, Isabella, Jackson, Kalamazoo, Kent, Lapeer, Lenawee, Livingston, Manistee, Mecosta, Menominee, Midland, Oakland, Ottawa, Sagi- naw, Saint Clair, Saint Joseph, Sanilac, Shiawassee, Van Buren, Washtenaw, and Wayne counties.

Genus PARABROCHYMENA Larivière, 1992 Parabrochymena arborea (Say), 1825. – Townsend (1890) reported this rough stink bug, a name commonly used for members of the genus, from Michigan (as Brochymena arborea). It has been taken under leaves at the edge of woodlands and on red maple (Acer rubrum L.) in Livingston County. Although primarily an herbivorous species, Hart (1919) recorded it attacking larvae of the Colorado potato beetle (Leptinotarsa decimlineata (Say), 1824), Larivière (1990, 1992, 1994) described and revised the new genus for the members of the arborea species group, and several studies done on genitalic morphology (Ahmad and McPherson 1998; McPherson and Ahmad 2005, 2007) support the monophyly of the group. Cuda and McPherson (1976) provided biological notes for this species. McDonald (1966) described the genitalia. Ruckes (1946) keyed the genus, and Larivière (1994) keyed the members of the genus. 32 specimens examined. Collection dates from 26 March to 1 November. Distribution: Barry, Clinton, Ingham, Ionia, Isabella, Kalamazoo, Lake, Lapeer, Livingston, Midland, Oakland, Ottawa, Saint Joseph, Shiawassee, and Washtenaw counties.

Tribe PENTATOMINI Leach, 1815 Genus AELIA Fabricius, 1803 Aelia americana Dallas, 1851. – McPherson (1970) reported this species from Michigan. Old World species of Aelia are well known pests of wheat (Kre- tovich 1944), and Panizzi et al. (2000) treated this as one of the less economically important species. McDonald (1966) described the genitalia. 5 specimens examined. Collection dates from 16 July to 7 October. Distribution: Cheboygan, Houghton, Luce, and Marquette counties. 2012 THE GREAT LAKES ENTOMOLOGIST 291 Genus BANASA Stål, 1860 Banasa calva (Say), 1832. – McPherson (1970) reported this species from Michigan. It has been taken at lights in Mason and Otsego counties. DeCoursey (1963) provided notes on the life history of this species. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. Thomas and Yonke (1981, 1985) reviewed, keyed, and provided a cladistic analysis of the genus, and Hoffman (2005) keyed several of the eastern species. 46 specimens examined. Collection dates from 27 May to 13 November. Distribution: Benzie, Ingham, Livingston, Mason, Otsego, Ottawa, Saint Joseph, and Washtenaw counties. Banasa dimidiata (Say), 1832. – This species was reported from Michi- gan by Stoner (1922) and Hussey (1922a). Stoner (1922) indicated this species showed a strong preference for white cedar (Thuja occidentalis L.) as a host, but a few specimens also were taken from round-leaved dogwood (Cornus rugosa Lam.), blueberry (Vaccinium sp.), and Canadian service berry (Amelanchier canadensis (L.) Medik.); it also was found in beach drift. This species has been taken on Drummond Island (Chippewa County) and Mackinac Island (Mackinac County). Label data indicate B. dimidiata has been taken in a hickory-oak forest in Calhoun County and at white and ultraviolet lights in Clinton, Livingston, Saint Joseph, and Washtenaw counties. This species has been taken on goldenrod (Solidago sp.) in Clinton County and chokecherry (Prunus virginiana L.) in Washtenaw County, as well as white cedar (Thuja occidentalis L.), and flowers of oxeye daisy Leucanthemum( vulgare Lam.) and carolina rose (Rosa carolina L.) in Cheboygan County. Over the last century, this pentatomid has been recorded variously under the original spelling, B. dimidiata (i.e., Froeschner 1988c) and the emendation B. dimidiata (i.e., Thomas and Yonke 1981, Hoffman 2005). DeCoursey (1963) provided notes on the life history of this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Thomas and Yonke (1981, 1985) reviewed, keyed, and provided a cladistic analysis of the genus, and Hoffman (2005) keyed several of the eastern species. 410 specimens examined. Collection dates from 20 April to 10 November. Distribution: Alcona, Alpena, Antrim, Arenac, Bay, Benzie, Berrien, Cal- houn, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Emmet, Genesee, Gladwin, Grand Traverse, Houghton, Huron, Ing- ham, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lapeer, Leelanau, Livingston, Mackinac, Macomb, Manistee, Marquette, Mason, Me- costa, Midland, Missaukee, Montcalm, Newaygo, Oakland, Oceana, Ontonagon, Osceola, Otsego, Presque Isle, Roscommon, Saginaw, Saint Joseph, Schoolcraft, Shiawassee, Tuscola, Van Buren, Washtenaw, Wayne, and Wexford counties. Banasa euchlora Stål, 1872. – Commonly called the juniper stink bug, one individual of B. euchlora, bearing the following label data, was examined by the author: MICHIGAN: Washtenaw Co., Ann Arbor, 10 January 1976, I. J. Cantrall, det. I. J. Cantrall 1976 [1 female] (UMMZ). A secondary label seems to negate the possibility of a mislabelled specimen: “Introduced? Found on cedar twigs in planter from local greenhouse.” It seems very likely that Cantrall’s suspicions were correct, and this individual represents an accidental introduction. This species is currently recorded from Alabama, Arizona, Arkansas, Colorado, Florida, Georgia, Idaho, Illinois, Indiana, Iowa, Maryland, Nevada, New Jersey, New Mexico, New York, North Carolina, Oklahoma, South Carolina, Texas, Utah, and Virginia (Froeschner 1988c) and shows a preference for juniper (Juniperus sp.) (Thomas and Yonke 1981). It should, however, be excluded from the Michigan faunal list, until it can be demostrated that a population persists in the state; nevertheless, it remains a potentially adventive species. The species is easily separated from other Michigan Banasa by the distinct white callosities at the basal angles of the scutellum. 292 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Banasa sordida (Uhler), 1871. – (NEW STATE RECORD). Two individuals of this pentatomid were examined by the author. Label data as follows: MICHIGAN: Saint Joseph Co., Klinger Lake, 9 May 1979, D. C. L. & N. M. Gosling, det. D. R. Swanson 2012 [1 male]; idem. 17 July 1983 [1 female] (UMMZ). The dates of the two collections indicate the likelihood of an established population in southern Michigan. Currently recorded from Arizona, California, Colorado, District of Columbia, Illinois, Massachusetts, Maryland, New Jersey, New Mexico, Utah, Virginia, Washington, British Columbia, and Ontario (Froe- schner 1988c), it is plausible that this species occurs in Michigan; furthermore, Thomas and Yonke (1981) characterized the distribution of B. sordida as “Entire U.S., northern Mexico and southern Canada.” The long rostrum, which reaches the base of the tubercle-bearing sternite, is a distinctive feature. Thomas and Yonke (1981, 1985) reviewed, keyed, and provided a cladistic analysis of the genus, and Hoffman (2005) keyed several of the eastern species. 2 specimens examined. Collection dates from 9 May to 17 July. Distribution: Saint Joseph County.

Genus CHINAVIA Orian, 1965 Chinavia hilaris (Say), 1832. – Hussey (1922a) reported the green stink bug from Michigan, where many individuals were collected from beach drift as well as beaten from various trees. A female sand wasp, Bicyrtes quadrifasciatus (Say), 1824 (Crabronidae: Bembicinae), was taken with a late-instar nymph of this pentatomid as prey in Oakland County. Label data indicate C. hilaris has been taken from a Malaise trap at 6 feet height in Newaygo County and white and ultraviolet lights in Livingston and Saint Joseph counties. This conspicuous species is known to attack several commercial crops and may cause significant damage in large numbers; Panizzi et al. (2000) dealt with the economic importance of this species. Olsen (1912) recorded this species occasionally attacking soft-boded larvae. McPherson and Tecic (1997) reported on the life history of this species. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. Rolston (1983) revised the group encompassing those species found in the United States. The members of this genus previously were placed in Acrosternum Fieber, 1860, with Chinavia variously accorded generic, subgeneric, or synonymic rank. However, Acrosternum has been restricted to a group of Old World taxa and the generic status of Chinavia seems to be well supported and generally accepted (Ahmad 1996; Schwertner 2005; Schwertner and Grazia 2006, 2007; Genevcius et al. 2012). 285 specimens examined. Col- lection dates from 10 January to 11 December. Distribution: Allegan, Baraga, Barry, Bay, Benzie, Berrien, Branch, Cal- houn, Cass, Cheboygan, Clinton, Eaton, Gratiot, Hillsdale, Houghton, Ingham, Isabella, Jackson, Kalamazoo, Lapeer, Lenawee, Livingston, Mackinac, Macomb, Marquette, Mason, Menominee, Midland, Monroe, Montcalm, Muskegon, Ne- waygo, Oakland, Oceana, Ottawa, Saginaw, Saint Joseph, Shiawassee, Van Buren, Washtenaw, and Wayne counties. Chinavia pensylvanica (Gmelin), 1790. – This species was reported from Michigan by Hussey (1922a), who noted a single specimen “swept from low vegetation at the base of a dune.” McDonald (1966) described the genitalia. Rolston (1983) revised the group encompassing those species found in the United States. As mentioned above, this species previously was placed in Acrosternum, a genus now restricted to a group of Old World species. 25 specimens examined. Collection dates from 5 June to 4 October. Distribution: Berrien, Cass, Ingham, Lapeer, Livingston, Saint Joseph, and Van Buren counties. 2012 THE GREAT LAKES ENTOMOLOGIST 293 Genus CHLOROCHROA Stål, 1872 Subgenus CHLOROCHROA Stål, 1872 Chlorochroa persimilis Horvath, 1908. – Townsend’s record (1890) of a nymph of Lioderma ligata is referred here. Stoner (1922) reported this species (as Chlorochroa uhleri) on smooth sumac (Rhus glabra L.), lowbush blueberry (Vaccinium angustifolium Aiton), and black huckleberry (Gaylussacia baccata (Wangenh.) Koch). Hussey (1922a) noted this species (under the same binomial as Stoner) taken on flowering spurge (Euphorbia corollata L.) on the dunes and among beach drift; nymphs also were frequently taken on milkweed (Asclepias sp.). Label data indicate this species has been taken on wild rose (Rosa blanda Gray) and sumac (Rhus sp.) in Cheboygan County as well as on common juniper (Juniperus communis L.) in Livingston County. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. Buxton et al. (1983) revised the sayi group, the complex to which this taxon belongs, and Thomas (1983) treated the group relationships within the genus. 156 specimens examined. Collection dates from 9 April to 17 November. Distribution: Alger, Baraga, Benzie, Berrien, Charlevoix, Cheboygan, Chippewa, Crawford, Emmet, Gladwin, Grand Traverse, Huron, Ingham, Iosco, Jackson, Kalkaska, Keweenaw, Lake, Lapeer, Livingston, Mackinac, Marquette, Mecosta, Midland, Missaukee, Montmorency, Muskegon, Oakland, Oscoda, Roscommon, Saint Joseph, Schoolcraft, Shiawassee, Washtenaw, and Wexford counties.

Genus COENUS Dallas, 1851 Coenus delius (Say), 1832. – This species was reported from Michigan by Stoner (1922) and Hussey (1922a). Stoner (1922) took a few individuals among beach drift and swept several specimens from wild raspberry (Rubus sp.). Hussey (1922a) noted this uncommon species “taken from grass by a creek...[and] at the edge of a cranberry bog.” Label data indicate C. delius has been taken on common mullein (Verbascum thapsus L.) in Livingston County and “[s]weeping brackensweet fern fallow field” in Isabella County. Oetting and Yonke (1971) provided biological information for this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Rider (1995) reviewed the genus. 139 specimens examined. Collection dates from 20 March to 9 November. Distribution: Allegan, Barry, Bay, Berrien, Calhoun, Cass, Cheboygan, Crawford, Emmet, Gogebic, Hillsdale, Huron, Ingham, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Lenawee, Livingston, Mackinac, Ma- son, Mecosta, Midland, Missaukee, Montmorency, Newaygo, Oakland, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Saint Clair, Schoolcraft, Van Buren, Washtenaw, and Wayne counties.

Genus COSMOPEPLA Stål, 1867 Cosmopepla lintneriana Kirkaldy, 1909. – This species was reported from Michigan by Townsend (1890) (as Cosmopepla carnifex) who reported “numbers flying in late summer,” although Hussey (1922a) found it “surprisingly uncommon” in Berrien County. Stoner (1922) swept this species from rough cinquefoil (Potentilla norvegica L.) in low moist areas. This species was taken on swamp milkweed (Asclepias incarnata L.) in Livingston County, parsnip (Pastinaca sativa L.) in Washtenaw County, and “feeding on Ranunculus seeds” in Che- boygan County. It also was collected from Stone Ring Marsh in the Edwin S. George Reserve (Livingston County). This species also has been collected on Isle Royale (Keweenaw County). In the past, C. lintneriana has been treated under the name Cosmopepla bimaculata (Thomas), 1865, but Rider and Rolston 294 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

(1995) clarified the nomenclature. Olsen (1910) and Fish and Alcock (1973) discussed its behavior. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. McDonald (1968b), McPherson (1976b), and McPherson and Tecic (1997) provided notes on the life history, and McDonald (1986) revised the genus. 718 specimens examined. Collection dates from 1 March to 13 November. Distribution: Alcona, Alger, Allegan, Baraga, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clinton, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Gogebic, Grand Traverse, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Lenawee, Livingston, Mackinac, Macomb, Marquette, Mason, Menominee, Midland, Missaukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Otsego, Presque Isle, Saginaw, Saint Clair, Saint Joseph, Sanilac, Schoolcraft, Shiawassee, Tuscola, Van Buren, Washtenaw, Wayne, and Wexford counties.

Genus DENDROCORIS Bergroth, 1891 Dendrocoris humeralis (Uhler), 1877. – McPherson (1970) reported this species from Michigan. This pentatomid has been taken in a prairie in Saint Joseph County and on black oak (Quercus velutina Lamb.) in Livingston County. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Nelson (1955) revised and Thomas and Brailovsky (1999) reviewed and keyed the genus. 62 specimens examined. Collection dates from 26 April to 13 November. Distribution: Berrien, Huron, Ingham, Jackson, Lake, Livingston, Mid- land, Newaygo, Oakland, Saint Joseph, Shiawassee, Van Buren, Washtenaw, and Wayne counties.

Genus EUSCHISTUS Dallas, 1851 Euschistus ictericus (Linnaeus), 1763. – This species was reported from Michigan by Hussey (1922a) who found it “sweeping sedges and grasses in more or less marshy localities, a few were secured from Saururus cernuus.” It also has been taken in a grass prairie in Calhoun County and from “[m]ixed iris-carex” in Livingston County. McPherson and Paskewitz (1984a) provided information on the life history of this species. Drickamer and McPherson (1992) compared the mating behavior of this species with several congeners and Thyanta custator accerra. 62 specimens examined. Collection dates from 17 May to 13 November. Distribution: Barry, Bay, Berrien, Calhoun, Cheboygan, Huron, Iosco, Kent, Lapeer, Livingston, Monroe, Oakland, Oceana, Saint Joseph, Washtenaw, and Wexford counties. Euschistus politus Uhler, 1897. – Blatchley (1926) included Michigan in the range of E. politus without citing specific localities. It has been collected from a Malaise trap in Lake County. McPherson (1974a) provided biological information for this species. Drickamer and McPherson (1992) compared the mating behavior of this species with several congeners and Thyanta custator accerra. 32 specimens examined. Collection dates from 11 May to 26 August. Distribution: Clare, Crawford, Huron, Iosco, Lake, Livingston, Manistee, Mecosta, Midland, Muskegon, Saint Joseph, Van Buren, and Washtenaw counties. Euschistus servus euschistoides (Vollenhoven), 1868. – This species was reported from Michigan by Townsend (1890) (as Euschistus fissilis). Stoner (1922) reported it from Michigan (as Euschistus euschistoides) and suggested it had a bivoltine life cycle around Douglas Lake, based on the emergence of nymphs. Stoner also recorded the species on smooth sumac (Rhus glabra L.), 2012 THE GREAT LAKES ENTOMOLOGIST 295 raspberry, and blackberry (Rubus spp.) as well as from open woods, grassy fields, cultivated areas, burned-over regions, and beach drift. Hussey (1922a) also took it from grassy areas, “especially in the hay fields and by the roadsides... and, more rarely, in clearings on the dunes”; he also found it abundant in beach drift. This species has been taken in from a field in Calhoun County, a moist meadow in Newaygo County, and Stone Ring Marsh in the Edwin S. George Reserve (Livingston County). Label data describe one locality as “Sweeping fallow roadside. Solidago dominant. Some shrubs.” in Isabella County. It has been collected from milkweed (Asclepias sp.) in Cheboygan County and common mullein (Verbascum thapsus L.) in Isabella and Livingston counties. A pair also was taken in copula on common milkweed (Asclepias syriaca L.) in Livingston County. This is, by far, the more common form of E. servus found in Michigan, although there is certainly some question as to the validity of the subspecies. In absence of definitive studies, I have chosen to conserve the traditional view of the two subspecies for this treatment. Youther and McPherson (1975) and Munyaneza and McPherson (1994) provided information on the life history of this species. Culliney (1985) summarized the occasional predaceous habits of this species. Panizzi et al. (2000) dealt with the economic importance of this species. Drickamer and McPherson (1992) compared the mating behavior of this species with several congeners and Thyanta custator accerra. Davidová- Vilímová and McPherson (1991) described and illustrated the male genitalia. Rolston (1974) revised the Euschistus species of Middle America. 553 specimens examined. Collection dates from 15 April to 13 November. Distribution: Alcona, Alger, Allegan, Arenac, Baraga, Barry,Bay, Berrien, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Dickinson, Eaton, Genesee, Gladwin, Gogebic, Grand Traverse, Gratiot, Hills- dale, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kent, Keweenaw, Lake, Lapeer, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Saint Joseph, Schoolcraft, Shiawassee, Van Buren, Washtenaw, Wayne, and Wexford counties. Euschistus servus servus (Say), 1832. – As Townsend (1890) had already reported the other subspecies as E. fissilis (see above), it is assumed that his record of E. servus refers to the nominate subspecies. Townsend reported it on the bud of a dandelion (Taraxacum officinale F. H. Wigg). Although the subspecies have been retained, all literature relevant to E. servus is listed under the above account. 6 specimens examined. Collection dates from 22 April to 18 August. Distribution: Cass, Clinton, Hillsdale, Ingham, Kalamazoo, and Saint Joseph counties. Euschistus tristigmus luridus Dallas, 1851. – Townsend’s (1890) record of E. tristimus is reported under this subspecies as it is by the far the more common form in Michigan. Still it should be noted that since his collections were taken in the southernmost line of counties in Michigan, this is very close to the purported overlap zone between the subspecies, and the record’s placement here is purely subjective. Townsend collected this species on holly-hock seed-cups (Alcea sp.). Stoner’s (1922) record of E. tristigmus also is referred here, and he reported the species from beach drift and such plants as smooth sumac (Rhus glabra L.), willow (Salix sp.), raspberry (Rubus idaeus L.), and blackberry (Rubus sp.). Hussey (1922a) reported of this species: “Fairly common...especially in damp localities; specimens were taken from Saururus cernuus, from marsh grasses, and from Benzoin aestivale...from wild rose and from the button-bush (Cephalanthus occidentalis). No specimens were taken in the dune region, nor were any found in the beach drift.” 296 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Label data indicate collections of E. t. luridus from many situations: a beach in Cheboygan County, a moist meadow in Newaygo County, night lights in Liv- ingston County as well as in open woods and strangely, under bark in Washtenaw County. One collection from Isabella County is described as “Sweeping fallow roadside. Solidago dominant. Some shrubs.” The species also has been taken from tamarack (Larix laricina (Du Roi) Koch), swamp birch (Betula pumila L.), and the leaves of common milkweed (Asclepias syriaca L.) in Livingston County as well as feeding on flowering spurge Euphorbia( corollata L.) in Isabella County and in copula on roadside elderberry (Sambucus sp.) in Washtenaw County. It has also been taken on flowers of several plants: meadow sweet Spiraea( alba Du Roi) in Montcalm County, sumac (Rhus sp.) in Benzie County, and oxeye daisy (Leucanthemum vulgare Lam.) in Cheboygan County. It also has been collected on Isle Royale (Keweenaw County). A large series of studies (McPherson 1971b, 1974b, 1975a, 1975b, 1976a, 1979a, 1979c; McPherson and Vangeison 1975; Clair and McPherson 1981; McPherson and Paskewitz 1982b) have sought to elucidate the life cycle and the relationship between certain variable morphological features and photoperiod in E. tristigmus. Additionally, the status of the subspecies remains questionable, although I have followed the traditional treatment. Culliney (1985) summarized the occasional predaceous habits of this species. Panizzi et al. (2000) treated this as one of the less economically important species. Drickamer and McPherson (1992) compared the mating behavior of this species with several congeners and Thyanta custator accerra. McDonald (1966) described the genitalia. Rolston (1974) revised the Euschistus species of Middle America. 866 specimens examined. Collection dates from 14 January to 21 December. Distribution: Alger, Allegan, Alpena, Antrim, Arenac, Baraga, Barry, Bay, Benzie, Berrien, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Goge- bic, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Presque Isle, Roscommon, Saginaw, Saint Clair, Saint Joseph, Sanilac, Schoolcraft, Shiawas- see, Tuscola, Van Buren, Washtenaw, Wayne, and Wexford counties. Euschistus tristigmus tristigmus (Say), 1832. – I could not locate the MSUC specimens referred to Euschistus tristigmus pyrrhocerus (Herrich-Schaeffer), 1841 by McPherson (1970). Furthermore, McPherson (1980a) subsequently did not include Michigan in the distribution of the nominate subspecies. Although the subspecies have been retained, all literature relevant to E. tristigmus is listed under the above account. No specimens examined. Collection dates (from McPherson 1970) from 18 July to 19 September. Distribution: Berrien, Hillsdale, and Kalamazoo counties. Euschistus variolarius (Palisot), 1817. – This species was reported from Michigan by Townsend (1890) from mullein in fields and raspberry bushes in gardens. Stoner (1922) recorded this species from lakeshore grasses and beach drift. Hussey (1922a) noted: “The most abundant pentatomid in southern Michigan...though less common in the dunes than in the other collecting localities. It occurred chiefly in the hayfields and on golden-rod, ragweed, and mullein; and during the second half of July it was the most abundant hemipteron found in the beach drift.” Label data indicate E. variolarius has been taken from red clover (Tri- folium pratense L.) in Branch, Calhoun, Eaton, Hillsdale, Ingham, Ionia, Kal- amazoo, Lenawee, Mecosta, Montcalm, Saint Joseph, and Washtenaw counties. Additional label data indicate it has been taken from beach drift in Berrien 2012 THE GREAT LAKES ENTOMOLOGIST 297 County, a farm field in Calhoun County, a moist meadow in Newaygo County, and in Stone Ring Marsh in the Edwin S. George Reserve (Livingston County). It has been taken on flowering spurge (Euphorbia corollata L.) and cultivated strawberries (Fragaria sp.) in Berrien County as well as willow (Salix sp.) in Huron County. This species has been collected in a rotary trap in a wheat field in Berrien County. Munyaneza and McPherson (1994) provided information on the life history of this species. Culliney (1985) summarized the occasional predaceous habits of this species. Panizzi et al. (2000) treated this as one of the less economically important species. Drickamer and McPherson (1992) compared the mating behavior of this species with several congeners and Thyanta custator accerra. Rolston (1974) revised the Euschistus species of Middle America. 744 specimens examined. Collection dates from 14 January to 28 November. Distribution: Alger, Allegan, Baraga, Barry, Bay, Berrien, Branch, Cal- houn, Cass, Cheboygan, Clinton, Crawford, Dickinson, Eaton, Emmet, Genesee, Gogebic, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Jackson, Kal- amazoo, Kent, Lake, Lapeer, Lenawee, Livingston, Macomb, Manistee, Mason, Mecosta, Midland, Missaukee, Monroe, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ottawa, Presque Isle, Roscommon, Saginaw, Saint Joseph, Schoolcraft, Shiawassee, Tuscola, Van Buren, Washtenaw, and Wayne counties.

Genus HALYOMORPHA Mayr, 1864 Halyomorpha halys (Stål), 1855. – Michigan was recently added to the list of states encompassing the growing range of the brown marmorated stink bug (Michigan Department of Agriculture and Rural Development 2011). However, this invasive Asiatic species is probably more widely distributed in the state than the sparse records indicate. Considered to be a major pest of many commercial crops, H. halys also may be a frequent and conspicuous overwintering guest in human habitations. Hoebeke and Carter (2003) reported this species from the United States, and Nielsen and Hamilton (2009) studied the life history of this species in the northeastern United States. 2 specimens examined. Collection dates from 30 April to 24 July. Distribution: Berrien, Eaton, Oceana, and Washtenaw counties.

Genus HOLCOSTETHUS Fieber, 1860 Holcostethus fulvipes (Ruckes), 1957. – McPherson (1979b) reported this species from Michigan, citing a specimen taken on common juniper (Juniperus communis L.) in Livingston County. McPherson also clarified that the specimen on which the Michigan records of Holcostethus abbreviatus Uhler, 1872 were based in his previous treatment (McPherson 1970) were misidentified individuals of H. fulvipes; thus, H. abbreviatus is not known from Michigan. McDonald (1974, 1982) revised the genus. 5 specimens examined. Collection dates from 12 May to 27 July. Distribution: Charlevoix, Livingston, and Montcalm counties. Holcostethus limbolarius (Stål), 1872. – This species was reported from Michigan by Townsend (1890) (as Peribalus limbolarius). Stoner (1922) reported a single individual from an oat field. Hussey (1922a) reported it abundant in the fields. Label data indicateH. limbolarius has been taken from a field in Calhoun County, corroborating the two authors’ observations. It has been collected on common mullein (Verbascum thapsus L.) and flowers of catsfoot Antennaria( sp.) in Livingston County as well as on cedar (Thuja sp.) and goldenrod (Solidago sp.) in Washtenaw County. This species also has been collected on Isle Royale (Keweenaw County). McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. McDonald (1974, 1982) revised the genus. 159 specimens examined. Collection dates from 13 April to 13 November. 298 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Distribution: Allegan, Bay, Benzie, Berrien, Calhoun, Cass, Cheboygan, Clinton, Crawford, Emmet, Gratiot, Huron, Ingham, Ionia, Jackson, Kent, Keweenaw, Lapeer, Livingston, Mackinac, Mecosta, Midland, Muskegon, Oak- land, Osceola, Ottawa, Saginaw, Saint Joseph, Schoolcraft, Tuscola, Van Buren, Washtenaw, and Wayne counties. Holcostethus mcdonaldi Rider and Rolston, 1995. – Furth (1974) reported this species from Michigan, and McPherson (1979b) corroborated the record and provided an additional locality. In the past, this species has been treated under the binomial Holcostethus piceus (Dallas), 1851, but Rider and Rolston (1995) clarified the status ofPentatoma picea and renamed the Nearctic taxon. McDonald (1974, 1982) revised the genus. 2 specimens examined. Collection dates from 21 to 26 June. Distribution: Alger and Cheboygan counties.

Genus HYMENARCYS Amyot & Serville, 1843 Hymenarcys nervosa (Say), 1832. – This species was reported from Michi- gan by Hussey (1922a) who noted it taken from fresh herbage by a spring, a hayfield, and under the bark of a beech stump at the edge of the woods. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Rolston (1973) reviewed the genus. 17 specimens examined. Collection dates from 11 February to 13 November. Distribution: Berrien, Ingham, Livingston, and Washtenaw counties.

Genus MCPHERSONARCYS Thomas, 2012 Mcphersonarcys aequalis (Say), 1832. – This species was reported from Michigan by McPherson (1970) under the genus Hymenarcys. Rolston (1973) noted several differences between H. aequalis and its congeners, and Thomas (2012) subsequently erected a genus for the species. Esselbaugh (1947) provided biological information for this species. 5 specimens examined. Collection dates from 4 May to 3 October. Distribution: Clinton, Crawford, Kalamazoo, Livingston, Washtenaw, and Wayne counties.

Genus MENECLES Stål, 1867 Menecles insertus (Say), 1832. – Hussey (1922a) reported Menecles incertus [sic] from Michigan in beach drift. Label data indicate this species has been taken on red maple (Acer rubrum L.) in Mason County. The deep set head is a unique character among the Michigan pentatomids. Balduf (1945) provided bionomic notes for this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Rolston (1972) reviewed the genus. 130 specimens examined. Collection dates from 23 March to 19 December. Distribution: Antrim, Berrien, Cheboygan, Clinton, Ingham, Ionia, Isa- bella, Lake, Lapeer, Livingston, Mason, Mecosta, Missaukee, Oakland, Saint Joseph, and Washtenaw counties.

Genus MORMIDEA Amyot and Serville, 1843 Subgenus MELANOCHILA Stål, 1872 Mormidea lugens (Fabricius), 1775. – This species was reported from Michigan by Townsend (1890) on leaves of oak (Quercus sp.). Stoner (1922) took a single nymph from reeds, and Hussey (1922a) found the species common, particularly in shaded grasses, and collected it from raspberry (Rubus sp.), 2012 THE GREAT LAKES ENTOMOLOGIST 299 beds of lizard’s tail (Saururus cernuus L.), and “sedge zones in the marshes.” The author has swept M. lugens from low grass in open woodlands in Oakland and Washtenaw counties, thus supporting one of Hussey’s observations. This species also has been taken in Stone Ring Marsh in the Edwin S. George Re- serve (Livingston County). However, M. lugens seems to be most commonly associated with grasses and sedges (McPherson 1974a, Rolston 1978); thus, several of the aforementioned plant records may represent incidental encounters. McPherson (1974a) provided biological notes for this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Rolston (1978) revised the genus. 368 specimens examined. Collection dates from 2 May to 13 November. Distribution: Alcona, Allegan, Arenac, Bay, Berrien, Branch, Calhoun, Cass, Cheboygan, Clare, Clinton, Crawford, Dickinson, Eaton, Emmet, Genesee, Gratiot, Huron, Ingham, Ionia, Iosco, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Lenawee, Livingston, Manistee, Mecosta, Menominee, Midland, Missaukee, Monroe, Montmorency, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Roscommon, Saginaw, Saint Joseph, Sanilac, Shiawas- see, Tuscola, Van Buren, Washtenaw, and Wayne counties.

Genus MURGANTIA Stål, 1862 Murgantia histrionica (Hahn), 1834. – Furth (1974) listed the harlequin bug among the Michigan fauna based on specimens purportedly housed in the UMMZ, but McPherson (1970, 1979b) could not locate the specimens. They remain unfound. The species, however, is included in the key to facilitate identification of legitimate records. Nevertheless, it should be dropped from the Michigan faunal list until the specimens are found or additional material is collected.

Genus NEOTTIGLOSSA Kirby, 1837 Subgenus NEOTTIGLOSSA Kirby, 1837 Neottiglossa trilineata (Kirby), 1837. – This species was reported from Michigan by Hussey (1922b). Blatchley (1926) and Torre-Bueno (1939) also included Michigan in the range for this species. No additional data other than that given by McPherson (1970) is available regarding this species in Michigan. McDonald (1966) described the genitalia. Rider (1989) revised the genus. 1 specimen examined. Collection dates is 31 May. Distribution: Mackinac County. Neottiglossa undata (Say), 1832. – The record of Neottiglossa sulcifrons Stål, 1872 reported by Townsend (1890) surely refers to this species as suggested by Hussey (1921). Hussey (1922a) found the species uncommon, but Stoner (1922) reported it numerous on bluegrass (Poa sp.) and redtop (Agrostis sp.) in association with aspen (Populus sp.) and other moist places. Label data indicate N. undata has been taken under oak (Quercus sp.) leaves in Livingston County and swept from an open field in Oakland County. It also has been collected on Isle Royale (Keweenaw County). Rider (1989) revised the genus. 234 specimens examined. Collection dates from 18 April to 13 November. Distribution: Alcona, Alpena, Antrim, Arenac, Barry, Bay, Benzie, Berrien, Charlevoix, Cheboygan, Clare, Clinton, Crawford, Dickinson, Emmet, Genesee, Gladwin, Gogebic, Hillsdale, Huron, Ingham, Ionia, Iosco, Iron, Jackson, Kal- amazoo, Kalkaska, Kent, Keweenaw, Lapeer, Lenawee, Livingston, Mackinac, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montmorency, Newaygo, Oakland, Otsego, Roscommon, Saint Joseph, Schoolcraft, Shiawassee, Van Buren, Washtenaw, Wayne, and Wexford counties. 300 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Genus OEBALUS Stål, 1862 Oebalus pugnax pugnax (Fabricius), 1775. – Commonly known as the rice stink bug, this species is easily recognized by the anteriorly-directed humeral spines. Steyskal (1938) reported this species from Michigan; no additional data is available regarding this species in Michigan. This is one of the more agriculturally destructive species; Panizzi et al. (2000) dealt with the economic importance of this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Sailer (1944) revised the genus under the name Solubea Bergroth, 1891 and subsequently clarified the validity of the generic name (Sailer 1957). 3 specimens examined. Collection dates from 30 May to 20 September. Distribution: Berrien and Wayne counties.

Genus PRIONOSOMA Uhler, 1863 Prionosoma podopioides Uhler, 1863. – This rare pentatomid was reported from Michigan in McPherson’s faunal list (1970); no additional data is available regarding this species in Michigan. The notched humeral angles render P. podopioides instantly recognizable among the pentatomids of Michigan. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. 2 specimens examined. Collection date is 14 August. Distribution: Hillsdale County.

Genus THYANTA Stål, 1860 Thyanta calceata (Say), 1832. – This species was originally reported as Thyanta custator (Fabricius), 1803 in the original Michigan list (McPherson 1970), but was subsequently corrected (McPherson 1979b); thus, T. custator is not known from Michigan. Many aspects of the life history and laboratory rearing of T. calceata have been described (Oetting and Yonke 1971, McPherson, 1977a, Paskewitz and McPherson 1982), including studies of the effect of varying photoperiod on the development of this pentatomid (McPherson 1977b, 1978a, 1978b; McPherson and Paskewitz 1982a, McPherson et al. 1983). Rolston and McDonald (1984) revised the concept of the genus, and Ruckes (1957) keyed the species. Ueshima’s (1963) cytological study also elucidated the specific status of some members of this genus. Rider and Chapin (1992) revised the North American species of the genus. 1 specimen examined. Collection date is 16 May. Distribution: Wayne County. Thyanta custator accerra McAtee, 1919. – Hussey’s record (1922a) of Thyanta custator certainly refers to this species. He reported it “swept from ragweed...[and] in the June beach-drift.” McPherson (1970) included this species as Thyanta pallido-virens accerra, like many other authors of the time. Label data indicate T. c. accerra has been taken on roundhead bushclover (Lespedeza capitata Michx.) and common mullein (Verbascum thapsus L.) in Livingston County. It also has been taken in a farm field in Calhoun County. McPherson (1979d) studied the morphological effects of varying photoperiod during the development of this species. Panizzi et al. (2000) treated this as one of the less economically important species. Drickamer and McPherson (1992) compared the mating behavior of this species with several species of Euschistus. Davidová-Vilímová and McPherson (1991) described and illustrated the male genitalia. Rolston and McDonald (1984) revised the concept of the genus in transferring several western species to the western Tepa Rolston and McDonald, 1984, and Ruckes (1957) keyed the species currently in Thyanta. Ueshima’s (1963) cytological study also elucidated the specific status of some members of this genus. Rider and Chapin (1992) revised the 2012 THE GREAT LAKES ENTOMOLOGIST 301 North American species of the genus. 136 specimens examined. Collection dates from 3 March to 9 November. Distribution: Antrim, Barry, Berrien, Branch, Calhoun, Cass, Clinton, Crawford, Grand Traverse, Ingham, Ionia, Jackson, Kalamazoo, Kent, Lapeer, Lenawee, Livingston, Mason, Montcalm, Newaygo, Oakland, Saginaw, Saint Joseph, Shiawassee, Van Buren, Washtenaw, and Wayne counties.

Genus TRICHOPEPLA Stål, 1867 Trichopepla atricornis Stål, 1872. – McPherson (1970) added T. atricornis to the faunal list of Michigan. The conspicuous pubescence of the dorsum renders species of this genus instantly recognizable among the state fauna, although this feature is shared with Prionosoma podopioides. McDonald (1976) revised the genus but did not include Michigan in the distribution of this species. 11 specimens examined. Collection dates from 11 June to 14 September. Distribution: Kalamazoo, Livingston, and Saint Joseph counties. Trichopepla semivittata (Say), 1832. – This species was reported from Michigan by Hussey (1922a), who found them “in the fields about the Warren Woods.” The conspicuous pubescence of the dorsum renders species of this genus instantly recognizable among the fauna of Michigan, although this feature is shared with Prionosoma podopioides. Label data indicate T. semivittata has been taken in a farm field in Calhoun County. McPherson (1972b) described the laboratory rearing of this species. McDonald (1966) described the genitalia of both sexes of T. semivittata; he later revised the genus (1976). Davidová- Vilímová and McPherson (1991) described and illustrated the male genitalia. 54 specimens examined. Collection dates from 3 March to October [no date]. Distribution: Barry, Bay, Berrien, Branch, Calhoun, Cass, Cheboygan, Clinton, Hillsdale, Ingham, Kalamazoo, Kent, Lapeer, Livingston, Mason, Midland, Oceana, Shiawassee, Van Buren, Washtenaw, and Wayne counties.

Tribe SCIOCORINI Amyot & Serville, 1843 Genus SCIOCORIS Fallén, 1829 Sciocoris microphthalmus Flor, 1860. – Stoner (1920) reported this small species from Michigan, and Hussey (1921) provided a corroborative account, in which he stated “I have also taken this species at Douglas Lake, from roadside grasses in a lowland stand of arbor-vitae, balsam fir, birch, and aspen.” The small size combined with the flattened habitus renders this species instantly recognizable among the Michigan fauna. McDonald (1966) described the genitalia. 7 specimens examined. Collection dates from 31 May to 18 July. Distribution: Cheboygan and Mackinac counties.

Subfamily PODOPINAE Amyot & Serville, 1843 Genus AMAUROCHROUS Stål, 1872 Amaurochrous brevitylus Barber and Sailer, 1953. – This turtle bug was reported from Michigan by McPherson (1970). Barber and Sailer (1953) revised the members of the subfamily in North America, and Davidová-Vilímová and McPherson (1994) discussed the history of the higher classification of the subfamily. 3 specimens examined. Collection dates from 13 May to 22 June. Distribution: Kalamazoo, Lenawee, and Mason counties. Amaurochrous cinctipes (Say), 1828. – Hussey (1922a) reported this species from Michigan (as Podops cinctipes) as “taken from Carex and Scirpus in the marsh.” This species also has been taken in sphagnum in late February in 302 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Washtenaw County. McPherson and Paskewitz (1984b) provided notes on the laboratory rearing and immatures stages of this species. McDonald (1966) and Davidová-Vilímová and McPherson (1991) described the genitalia. Barber and Sailer (1953) revised the turtle bugs of North America, and Davidová-Vilímová and McPherson (1994) discussed the history of the higher classification of the subfamily. 58 specimens examined. Collection dates from 23 February to 13 November. Distribution: Bay, Berrien, Branch, Cheboygan, Clinton, Gladwin, Ingham, Kalamazoo, Kent, Livingston, Mason, Mecosta, Midland, Oakland, Saint Joseph, Van Buren, Washtenaw, and Wayne counties.

Acknowledgments I am grateful to Mark O’Brien (UMMZ) and Gary Parsons (MSUC) for the privilege of studying the material under their care. I also thank Dr. David A. Rider, North Dakota State University, Fargo, North Dakota, for generously sharing his expertise, particularly involving unresolved taxonomic issues. I owe much appreciation to two anonymous reviewers, whose insightful comments greatly improved the manuscript.

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Abstract The Pentatomoidea fauna for North Dakota is documented. There are 62 species of Pentatomoidea known from North Dakota: Acanthosomatidae (2), Cydnidae (4), Pentatomidae: Asopinae (9), Pentatomidae: Pentatominae (34), Pentatomidae: Podopinae (2), Scutelleridae (6), and Thyreocoridae (5). Of this total, 36 represent new state records for North Dakota. Additionally, 16 new state records are reported for Minnesota, and one new state record each for South Dakota, Texas, and Utah. The new state records for North Dakota are: Acantho- somatidae: Elasmostethus cruciatus (Say), Elasmucha lateralis (Say); Cydnidae: Amnestus pusillus Uhler, Amnestus spinifrons (Say), Microporus obliquus Uhler; Pentatomidae (Asopinae): Perillus exaptus (Say), Podisus brevispinus Phillips, Podisus maculiventris (Say), Podisus placidus Uhler, Podisus serieventris Uhler; Pentatomidae (Pentatominae): Aelia americana Dallas, Neottiglossa sulcifrons Stål, Euschistus ictericus (Linnaeus), Euschistus latimarginatus Zimmer, Euschistus variolarius (Palisot de Beauvois), Holcostethus macdonaldi Rider and Rolston, Menecles insertus (Say), Mormidea lugens (Fabricius), Trichope- pla atricornis Stål, Parabrochymena arborea (Say), Mecidea minor Ruckes, Chinavia hilaris (Say), Chlorochroa belfragii (Stål), Chlorochroa ligata (Say), Chlorochroa viridicata (Walker), Tepa brevis (Van Duzee), Banasa euchlora Stål, Murgantia histrionica (Hahn); Pentatomidae (Podopinae): Amaurochrous brevitylus Barber and Sailer, Amaurochrous cinctipes (Say); Scutelleridae: Phimodera binotata (Say), Vanduzeeina borealis Van Duzee, Acantholomidea denticulata (Stål), Homaemus bijugis Uhler; and Thyreocoridae: Corimelaena lateralis (Fabricius), Galgupha ovalis Hussey. New state records for Minnesota are: Acanthosomatidae: E. cruciatus; Cydnidae: Amnestus pallidus Zimmer, A. pusillus, M. obliquus; Pentatomidae (Asopinae): P. placidus; Pentatomidae (Pentatominae): A. americana, N. sulcifrons, E. ictericus, E. latimarginatus, Holcostethus abbreviatus Uhler, Holcostethus limbolarius (Stål), M. insertus, M. lugens, T. atricornis, C. belfragii; and Scutelleridae: V. borealis. B. euchlora is recorded from South Dakota for the first time, andE. latimarginatus is reported from Texas and Utah for the first time. Keys are provided for the identification of all included taxa.

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Introduction The taxonomic literature is wrought with an abundance of faunal studies of various groups from various states, regions, and/or countries. The introductory notes of nearly every one of these elaborates on the great diversity of habitats contained in the study area to give cause for the study. At first glance, North Dakota may appear to be an exception; the perception being that North Dakota is composed wholly of grassland prairie. Although it is true that the majority of

1Department of Entomology, North Dakota State University, NDSU Dept. 7650, P. O. Box 6050, Fargo, ND 58108-6050. 2012 THE GREAT LAKES ENTOMOLOGIST 313 the state is covered with grassland prairie, upon closer inspection we begin to see that even North Dakota has a variety of habitats to survey. The grassland prairie itself includes areas of rolling sand dunes, especially in the southeastern counties, and ranges from mixed grass prairies with pockets of tallgrass in the east to widespread shortgrass prairie in the west. Many insect species usually considered to be of eastern origin enter North Dakota in the southeastern part of the state, often in the riparian habitats up and down the Red River Valley, but also along some of the more western rivers (Wild Rice, Sheyenne, etc.). Of course, there is a boreal component in the northern part of the state, especially in the Turtle Mountains region. The rolling sand dunes occur mainly in the Sheyenne National Grasslands (SNG), but there are a few isolated areas with dunes in central North Dakota; these possess faunas similar to dune areas in South Dakota and Nebraska. Finally, the southwestern areas of the state are much drier, the insect fauna containing species more indicative of the desert southwest. Nevertheless, the view that North Dakota was nearly void of interesting habitats to survey has led to a paucity of collecting in the state. I am aware of only two major collecting expeditions within the state. The U.S. Geological and Geographical surveys of the 1860’s and 1870’s included North Dakota. The results of these expeditions were reported on by the eminent hemipterist P. R. Uhler, but in his early work, he often gave the locality data as simply “Dakota” (e.g., Uhler 1872). Later, Uhler (1878) reported on the Hemiptera collected by Dr. Elliott Coues from a collecting expedition through North Dakota and into Montana; in this work he gave more specific locality details. The second important collecting expedition was made in 1920 by T. H. Hubbell as part of a joint venture of the newly formed (1919) North Dakota Biological Station and the Museum of Zoology of the University of Michigan. The results of this expedition were reported upon by another well-known hemipterist, R. F. Hussey (1922). Specimens from the Uhler studies may be conserved in the U.S. National Museum of Natural History in Washington, D.C.; specimens from the latter expedition are housed at the University of Michigan (Dan Swanson, personal communication). The only other North Dakota records of Heteroptera come from specific revisions of certain genera, and as such are scattered throughout the literature. There are several papers documenting the Heteroptera fauna of neighboring South Dakota (Parshley 1922; Harris 1937, 1943). There really is no good comprehensive list of terrestrial Heteroptera for Minnesota; the only records come from scattered revisions and lists from neighboring states, and a few minor records from an outdated, extension-style publication (Lugger 1900) The present contribution is the first part of a planned series of papers which will document the entire Heteroptera fauna of North Dakota. It is intended that all parts will include keys for the identification of species which are known from North Dakota (frequently the keys will also include non-Dakotan species that may eventually be found in the state). This part covers the Pentatomoidea. In recent years, the taxonomic staff at North Dakota State University has also conducted intensive collecting in western areas of Minnesota, which has provided several new heteropteran records for that state; when convenient those records are also reported upon within this series of papers. Other than scattered regional treatments or revisions of specific genera, the best sources for studying the eastern North American pentatomoid fauna remain Blatchley (1926) and McPherson (1982). The former treats all Heteroptera and covers the eastern U.S., but specializes mainly on the faunas of Indiana and Florida; the latter treats only the Pentatomoidea, and covers the northeastern United States with an emphasis on Illinois. McPherson (1982) provided many references to distribution, life history, food plants, natural enemies, most of which are not repeated here. I also find the “How to Know the True Bugs” by Slater and Baranowski (1978) immensely helpful in keying specimens to at least the generic level, and occasionally on to species. A more recent world 314 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 synthesis of the family classification has been presented by Schuh and Slater (1995). Specific references for various generic revisions will be given under the treatment of those genera. It is with great pleasure that this manuscript is dedicated to Dr. Jay McPherson upon the occasion of his retirement. Jay has trained a plethora of students in hemipteran taxonomy, biology, life history; but more importantly he has instilled an intense interest (or excitement) in these students for this wonderful group. Although I was not one of Jay’s students, he provided sage advice while I worked on my dissertation research, and has continued to give encouragement through the years. I have become good friends with a number of his students, all of whom received wonderful training from Jay. His legacy will continue through these students, and others (such as myself) that he has influenced through the years. Thanks Jay!

Materials and Methods The basis for this study (and for the future parts of this series) comes from both the literature and from collected specimens. The main literature sources were mentioned in the introduction, but it would be prudent to mention the main collecting sites listed in those papers. In some of Uhler’s earlier papers (e.g., 1872), he simply listed “Dakota” as the locality for certain species. These records probably originated from North Dakota as I have not seen any specific South Dakota records in any of Uhler’s papers. In his 1878 paper, Uhler included only four collection sites, one of which is in Montana. His sites included Pembina (Pembina Co.), Turtle Mountains (probably in Rollette Co.), Mouse River (probably in Bottineau Co.), and Milk River (Montana). The details of the Hubbell expedition are outlined nicely by Hussey (1922) (see that paper for details). In general, Hubbell collected in the following localities: 1) Devils Lake region (Ramsey Co.) with several short expeditions to Stump Lake and Spring Lake (Nelson Co.), and to a spot near the Sheyenne River (Eddy Co.); 2) Turtle Mountains (probably in Rollette Co.) with a few days in Bottineau (Bottineau Co.); 3) southwestern areas near Amidon (Slope Co.); and 4) Fargo (Cass Co.). Specimens from the following four collections were examined for this study (acronyms in parentheses). The bulk of the study material comes from the North Dakota State Insect Reference Collection (NDSU). This collection contains specimens from throughout the state, with some emphases on the southeastern and southwestern regions (also western areas of Minnesota). Other collections examined include those belonging to the University of North Dakota (UNDG) and Minot State University (MISU), which have emphasis on the faunas from Grand Forks Co. and Ward Co., respectively. The fourth collection studied is my own (DARC) composed of specimens from my years of collecting, trading, etc. To aid in future sampling, additional information about a few of the collecting sites is given here. Within the data listed in this paper, the reader will notice quite a few references to Mirror Pool and Mirror Pool WMA. There are two (maybe more) old oxbows of the Sheyenne River (in the Sheyenne National Grasslands in southeastern North Dakota, and abbreviated SNG throughout this work) that are known as Mirror Pool. The first, sometimes referred to as Iron Pond by locals, lies south of the river in Richland County. This is a beautiful U-shaped pond with the arms of the ‘U’ northward near the river, and a greatly elevated ridge following around the southern shore of the pond. This pond is officially a State Natural Area (SNA). Access to this pond was by a lightly maintained dirt road that was slowly eroded away in several places by the main river. This road was eventually closed, gated, and locked several miles away from the pond in each direction. The second Mirror Pool is actually a series of 2012 THE GREAT LAKES ENTOMOLOGIST 315 small to medium-sized ponds located north of the Sheyenne River in Ransom County, but still only a couple miles west of the Mirror Pool described above; this area is officially referred to as the Mirror Pool Wildlife Management Area (or WMA). Early access to this area was by a two-wheel track across several miles of pasture. Eventually, a new gravel road was constructed that has greatly helped access to this area. For our recent work, collecting switched from one site to the other roughly around the year 2000; references to Mirror Pool prior to 2000 generally refer to the site in Richland County; those after 2000 probably refer to the site in Ransom County. There are also many references to the Horsetrail Head site. This site is on County Road 23 in the eastern part of the SNG. County Road 23 is a gravel road that crosses the Sheyenne River and winds southward through rolling sand dunes populated with many trees, especially bur oak. This is why this area is affectionately called the Oak Savannah (at least by our personnel). Approximately two miles south of where the road crosses the Sheyenne River is a small grass parking lot on the east side of the road. This parking area is equipped with special log posts for tying up horses, and a horse trail winds off through the dunes both to the east and to the west. We have sampled on both sides of the road, hence the differences in longitude and latitude listings for this site. Another site commonly surveyed is the Ekre Grassland Preserve. This is a parcel of land that was donated to NDSU, and is presently under the supervi- sion of the Range Program. It is located on the east side of State Highway 18, approximately 5 miles south of State Highway 46. This site has a house and a few barns with pastures; there are some forested areas, and a fairly large tallgrass prairie habitat on the side of a large hill. The devastating floods we have experienced during the past 10-15 years have created several interesting new places to collect in our area. Because of repeated flooding, the Fargo city government has purchased (bought out) all the homes in several neighborhoods along the Red River. These homes have been torn down and the lots cleaned of debris. The entrance to these neighborhoods have usually been gated and locked, and the land has been allowed to return to its native habitat (most are in transition with a lot of weeds, etc.). This has resulted in areas with fairly native habitat right along the river with transitional habitats where the homes used to be, and there are still paved roads that allow for easy walking access. Examples of this type of habitat cited in this paper are the Chrisan Edition, Wild Rice River at University Drive, and Wild Rice River at Red River. The arrangement of taxa in this paper is purely alphabetical by family, subfamily, tribe, genus, and species. The tribal classification used in this paper is tentative at best. It is based primarily on the taxonomic literature available, and perhaps some on my own work. It should be noted, however, that a thorough phylogenetic study on the tribal classification of the Pentatomidae was not conducted, and the tribal arrangement used herein will invariably change once such a study is completed. It should also be noted that the keys contained within this paper should work for those species occurring in North Dakota, and perhaps adjoining areas. They will not work for other regions of the United States. The keys were developed with ease of use as the primary focus. No attempt was made to make the keys reflect phylogeny. The exception to this, out of some ne- cessity, may be the key to Pentatominae tribes. For this reason, I have also developed a key to the genera of all Pentatominae known to occur in North Dakota, irrespective of what tribe they belong. In some ways, this key may be easier to use than first keying the specimen to tribe, and then keying to genera within each tribe. At any rate, this will give the user two routes for identifying their specimens. 316 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

KEY TO NORTH DAKOTA FAMILIES OF PENTATOMOIDEA 1 Tarsi two-segmented...... Acanthosomatidae – Tarsi three-segmented...... 2 2(1) Hind tibiae provided with one to several rows of stout spines...... 3 – Hind tibiae lacking stout spines (hairs may be present, but they are never spine-like)...... 4 3(2) Scutellum subtriangular in shape...... Cydnidae – Scutellum large, rounded, covering most of abdomen...... Thyreocoridae 4(2) Scutellum greatly expanded and rounded, covering nearly all of hemelytra, sometimes (Eurygaster) more spatulate in shape, but in this case, the exposed portion of the corium narrows apically, apex angulate to acute...... Scutelleridae – Scutellum usually subtriangular, sometimes (Coenus) more spatulate in shape, but in this case, exposed portion of corium widens apically, apex rounded to truncate...... Pentatomidae

ACANTHOSOMATIDAE Signoret, 1863 This family is easily separated from all other North American pentatomoid families by having only two tarsal segments. All other North American pentatomoid families have three tarsal segments (note that there are several exceptions in other geographical regions). The scutellum is subtriangular, and the tibiae lack stout spines or bristles. The coloration is often brown to mottled gray, but may be paler, and may even have reddish markings. Thomas (1991) provided a review of the genera and species for North America. At present, only two genera are known to occur in North America, both of which are now known from North Dakota. Many species are known to provide parental care to eggs and early instars, and they often are found breeding on members of the Betulaceae. The Acanthosomatidae is divided into three subfamilies (Acantho- somatinae, Blaudusinae, Ditomotarsinae). Both our North American genera belong in the nominate subfamily.

Key to the genera of Acanthosomatidae in North Dakota 1 Ostiolar canal relatively short, extending laterally only to middle of metapleuron; posterolateral angles of pronotum prominent, distinctly produced posteriorly...... Elasmucha – Ostiolar canal relatively long, extending laterally beyond middle of metapleuron; posterolateral angles of pronotum angulate, but not prominent, not produced posteriorly...... Elasmostethus

Elasmostethus Fieber, 1860 The genus Elasmostethus is represented in North America by three species, one of which is now known from North Dakota. Elasmostethus interstinctus (Linnaeus) is a Palearctic species known in the New World only from Alaska; E. atricornis (Van Duzee) appears to be an eastern species, not definitely occurring further west than Illinois and Michigan. Torre-Bueno (1939) (repeated by McPherson 1982) listed Montana, but the source of this record is unknown, and is in need of verification. Elasmostethus cruciatus (Say) is herein reported from North Dakota for the first time. North American species ofElasmostethus tend to be somewhat larger in size than North American species of Elasmucha, and can be recognized by the longer ostiolar rugae. 2012 THE GREAT LAKES ENTOMOLOGIST 317

Elasmostethus cruciatus (Say, 1831) - For comparative purposes E. cruciatus has the antennae pale with segments four, five, and occasionally three darker, the humeral angles are pale, often tinged with red, and the pronotal punctures are dark. In E. atricornis, the antennae are entirely black, the humeral angles are black, and the pronotal punctures are concolorous with the pronotal surface. McPherson (1982) gave the distribution as Labrador and Québec west to the Pacific Coast, and south to South Carolina, northern Georgia, Texas, New Mexico, and California. Froeschner (1988a) did not list this species from North Dakota, or from any of the surrounding states or provinces, but Maw et al. (2000) recorded it from both Manitoba and Saskatchewan. Lugger (1900) listed this species from northern Minnesota, but his figure is clearly notE. cruciatus; the pronounced posterolateral pronotal angles, the darkened humeral angles, and the black marks along the connexival incisures indicate that he actually had Elasmucha lateralis. Jones and McPherson (1980) studied the biology of this species, and reported that in South Carolina, it feeds and reproduces on common alder, Alnus serrulata (Aiton) [Betulaceae]. Specimens Examined: MINNESOTA [New State Record]: Cass Co.: Gull Lake, 30-VIII-1996, Plant (1♀ DARC). St. Louis Co.: Giants Ridge, T059N R16W NE13, 16 May 1992, D. Cuthrell, light trap (1♂ NDSU). NORTH DAKOTA [New State Record]: Cass Co.: north Fargo, 17 Aug. 1999, P. Mason, on Betula (1♀ NDSU).

Elasmucha Stål, 1864 Elasmucha contains two U.S. species, one of which (cordillera Thomas) is known only from New Mexico southward; the other species, E. lateralis (Say), occurs across the northern U.S. and southern Canada, and is herein reported from North Dakota for the first time. North American species of Elasmucha tend to be smaller than North American species of Elasmostethus, and can be recognized by the shorter ostiolar rugae. Elasmucha lateralis (Say, 1831) - This species is smaller in size than Elasmostethus cruciatus, and the ostiolar ruga is shorter, not reaching the middle of the metapleuron. McPherson (1982) gave its distribution as Québec and New England west to the Pacific Coast, and south to South Carolina. Froeschner (1988a) listed this species from Minnesota, and Maw et al. (2000) recorded it from both Manitoba and Saskatchewan. Several specimens have been examined from northern Minnesota as well as the North Dakota specimens listed below. There are numerous papers citing birch [Betulaceae] as the host plant of this species (most recently, see Jones and McPherson 1980, who also studied its biology, and confirmed the presence of maternal care of the young). Specimens Examined: NORTH DAKOTA [New State Record]: Grand Forks Co.: Forest R. Biol. Area, 1.8 mi. W, 1 mi. N Inkster, T154N R55W, Sec. 11 & 14, 16 July 1977 (2♀♀ UNDG).

CYDNIDAE Billberg, 1820 Members of the family Cydnidae are relatively easy to recognize. They generally are small to medium in size, uniformly black, brown, or reddish-brown in color, have a triangular scutellum, and have one to several rows of stout spines on the hind tibiae. Their common name is burrowing bugs, and most occur in or on the ground; occasionally members of the Sehirinae are found on foliage. Many are collected at lights. The best reference for North American cydnids remains the monographic work of Froeschner (1960). Prior to this study, only one species had been reported from North Dakota; three more species in two genera are herein added. 318 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Key to the Subfamilies and Genera of North Dakota Cydnidae 1 Hemelytra with clavi meeting along midline posterior to apex of scutellum forming a straight line, the length of which is about one-fourth to one-third the length of the scutellum (Amnestinae)...... Amnestus – Hemelytral clavi not meeting along midline posterior to scutellum...... 2 2(1) Anterolateral margin of pronotum provided with a row of submargnal setigerous punctures; diameter of tarsal segment II subequal to diameter of tarsal segments I and III (Cydninae)...... Microporus – Anterolateral margin of pronotum lacking submarginal row of setigerous punctures; diameter of tarsal segment II distinctly less than diameter of tarsal segments I and III (Sehirinae)...... Sehirus

Amnestus Dallas, 1851 Amnestus together with Lattinestus Eger and a handful of fossil genera form the subfamily Amnestinae. This subfamily is easily recognized by the clavi meeting in a straight line posterior to the scutellar apex. Members of Amnestus tend to be quite small (less than 5mm in length), and are often uniformly reddish-brown in color (a few, including spinifrons, may be somewhat darker). Until recently, it was believed that Amnestus was entirely a New World genus, but a single species, A. pusillus Uhler has been discovered in Iran (Lis 1998, Martinez et al. 2012). There are six species known to occur in the United States, but two of these, pusio (Stål) and trimaculatus Froeschner, only occur in the U.S. from south Texas and Florida, respectively. Amnestus basidentatus Froeschner is more widespread in the southeastern U.S.; the closest record to North Dakota coming from Missouri. The remaining three species are all now known to occur in North Dakota, or nearby in western Minnesota; they can be identified by the following key.

Key to the Species of Amnestus in North Dakota 1 Juga each with four marginal pegs...... Amnestus pusillus Uhler – Juga each with five marginal pegs...... 2 2(1) Smaller, usually less than 2mm in length, coloration reddish brown; length of rostrum tends to be shorter, not reaching base of abdomen [not yet found in North Dakota]...... Amnestus pallidus Zimmer – Larger, usually more than 2mm in length; coloration dark brown to fuscus; length of rostrum tends to be longer, reaching nearly to or beyond base of abdomen...... Amnestus spinifrons (Say) Amnestus pallidus Zimmer, 1910 - Traditionally, the length of the rostrum has been used to separate this species from A. spinifrons, it being cited as longer in spinifrons. The length of the rostrum seems fairly stable in specimens of A. pallidus, but it seems to vary in A. spinifrons (see discussion below). My concept of this species is that it tends to be smaller, and more reddish-brown than A. spinifrons. McPherson (1982) gave the distribution for this species as Québec, Ontario, and Massachusetts south to Georgia, and west to Washington and California. Froeschner (1988b) did not list this species from North Dakota, or from any of the surrounding states or provinces. Specimens have been examined from western Minnesota, so it is likely that it will eventually be found in North Dakota. Not much is known about the biology of this species, but it has been collected on the aster, Antennaria plan- taginifolia (Linnaeus) (Stoner 1920). Specimens examined: MINNESOTA [New State Record]: Clay Co.: Fel- ton Prairie, 47° 03′Lat. 96° 26′Long., 6-X-1999, C. Davis & C. Jordan, pitfall trap 2012 THE GREAT LAKES ENTOMOLOGIST 319 (1♂, NDSU); CCL Trust Lands, 47° 03′Lat. 96° 26′Long., 20-V-1999, P. Tinerella & G. Fauske, pitfall trap (2♀♀, NDSU), 26-VI-1998, P. P. Tinerella, pitfall trap (1♀, NDSU). Amnestus pusillus Uhler, 1876 - This is the only species of Amnestus in our area that has only four marginal pegs on each jugum; both A. pallidus and A. spinifrons have five. This species has not previously been reported from North Dakota, or from any of the surrounding states or provinces. McPherson (1982) gave the distribution as Québec, Ontario, and Maine west to Oregon, and south to Virginia, Louisiana, Texas, California, and Mexico. Specimens Examined: MINNESOTA [New State Record]: Clay Co.: Bluestem Prairie, 46° 51′Lat. 96° 29′ Long., 10-VI-1999, P. Tinerella & G. Fauske (1♂ NDSU). NORTH DAKOTA [New State Record]: Cass Co.: Fargo, NDSU, 14-VI-2011, D. Rider & G. Fauske, Lindgren trap baited with manuka oil (1♀ NDSU), 8-IX-2010, D. Rider & G. Fauske, Lindgren trap baited with lineatin lure (1♂ NDSU). Sioux Co.: Fort Yates, 24 Aug. 1988, J. T. Schultz (1♂ NDSU). Amnestus spinifrons (Say, 1825) - This species, along with A. pallidus, has five marginal pegs on each jugum. Specimens of this species tend to be larger and much darker brown to fuscus in color than A. pallidus. Traditionally, this species has been described as having a much longer rostrum than A. pallidus. My concept of the species indicates that this might be true in more southern parts of its range, but specimens from northern areas tend to have the rostrum similar in size with those of A. pallidus. McPherson (1982) provided the following distribution for this species: Québec, Ontario, and Massachusetts south to Florida, and west to Iowa, Kansas, Texas, and Utah. This species has not previously been recorded from North Dakota, or any surrounding state or province. Specimens Examined: NORTH DAKOTA [New State Record]: Cass Co.: Fargo, NDSU, 14-VI-2011, D. Rider & G. Fauske, Lindgren trap (2♂♂ NDSU), 27-VI-2011, D. Rider, G. Fauske, Lindgren trap (1♂ 1♀ NDSU).

Microporus Uhler, 1872 Microporus is a member of the nominate subfamily, and can be separated from all other cydnine genera by the metapleural evaporative area that just outlines the peritreme. It is the only cydnine genus known to occur in North Dakota. Microporus obliquus Uhler, 1872 - This species is relatively easy to separate from other North Dakota cydnids; it is relatively small, it lacks the claval commissure of the Amnestinae, and it has many long hairs dorsally. Froeschner (1988b) listed this species from South Dakota, and Maw et al. (2000) recorded this species from Saskatchewan. McPherson (1982) gave the distribution for this species as Virginia, South Carolina, and Louisiana west to Washington, Oregon, California, and Mexico. It is interesting to note that this species has in the past been associated with sandy areas (Vestal 1915, Hart 1919, Stoner 1920); two of the collection sites listed below (Agassiz Dunes in western Minnesota, and Ransom County) are largely composed of sandy areas. Specimens Examined: MINNESOTA [New State Record]: Polk Co.: Agassiz Dunes, 47 30′Lat. 96 17′Long., 11-VI-1999, P. Tinerella & G. Fauske, pitfall trap (1♀ NDSU). NORTH DAKOTA [New State Record]: McHenry Co.: Sec. 36 Twp. 156 R. 78, 27-VII-1967, J. Knudson (1♂ NDSU). Ransom Co.: 29-IX-1960, J. Onsager (1♀ NDSU).

Sehirus Amyot and Serville, 1843 Sehirus is a member of the subfamily Sehirinae, which contains several genera and many Palearctic species. In the New World, the Sehirinae is rep- 320 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 resented by a single species, S. cinctus (Palisot de Beauvois) which has been divided into three subspecies: the southern texensis Froeschner, the widespread nominate form, and the northern albonotatus Dallas. This species is separable from all other North American cydnids by the white margins of the pronotum, corium, and hemelytra. Sehirus cinctus albonotatus Dallas, 1851 - This species also lacks the claval commissure found in the amnestines; it is somewhat larger than Micro- porus and it lacks the long dorsal hairs which characterize that species. This subspecies has a small white spot near the apex of R+M vein on each corium (these white spots are lacking in the nominate subspecies). McPherson (1982) gave the distribution of this species as Newfoundland and Maine south to Penn- sylvania, and west to Alberta and northern California. Uhler (1878) reported this species from two North Dakota localities: Bottineau Co.: near Mouse River, August 29, 1973, and Pembina Co.: near Pembina. All known records of this species come from the eastern half of North Dakota. Lugger (1900, as Canthophorus cinctus) recorded this species from Minnesota. Although other cydnids seem to remain close to or in the soil, this species can often be caught by sweeping various herbs, especially certain mint species (Lamiaceae). McDonald (1968a) reared this species in the laboratory on Stachys palustris Linnaeus [La- miaceae], and described the eggs and various nymphal instars. Five specimens in the NDSU collection were collected from Physostegia parviflora Nutt. ex A. Gray [Lamiaceae], and several more were collected at lights; the Fort Ransom specimens I collected (see below) were also on an unidentified mint. Specimens Examined: NORTH DAKOTA: Cass Co.: Fargo, 11-VI- 1924, R. L. Webster (1♀ NDSU), 5-IX-1919, O. A. Stevens, Physostegia parviflora (1♂ 2♀♀ 1? NDSU); NDSU Campus, 4-IX-2001, D. A. Rider (1♂ DARC); Wild Rice R. at Red Riv., 46.758N, 96.787W, 1-VIII-2012, D. Rider (1♂ DARC), 13-VIII-2012, D. A. Rider (13♂♂ 3♀♀ DARC); 18-VII-1960, D. Noetzel (1♀ NDSU), 21-VII-1971, D. Gardner (1♀ NDSU), 9-VIII-1968, J. Johnson (1♀ NDSU). Grand Forks Co.: Northwood, 15-VI-1964, R. L. Post (2♂♂ NDSU), 11-VII-1963, R. L. Post (6♂♂ NDSU). Griggs Co.: Cooperstown, 30-VII-1964, R. J. Saver (2♂♂ NDSU). LaMoure Co.: 22-VI-1959, R. L. Post (1♂ NDSU). Ransom Co.: Fort Ransom St. Pk., 13-VII-2002, D. A. Rider (10♂♂ 1♀ DARC); 2.5km. SW McLeod, 12-VII-1995, D. Rider, black light & mercury vapor light (4♀♀ DARC); 2mi. SW McLeod, T134N R53W SW35, 14-VII-1992, D. A. Rider (2♀♀ DARC); SNG Orchid Site D, T134N R53W SE 5, 27-VII-1992, D. Cuthrell (1♀ DARC).

PENTATOMIDAE Leach, 1815 The family Pentatomidae is largest of the pentatomoid families containing nearly 5,000 species. It has been divided into several subfamilies, five of which have members occurring in the United States. Of these five, two (Discocephali- nae, Edessinae) only have a few species each occurring in the very southern areas of the U.S. The remaining three (Asopinae, Podopinae, and the nominate subfamily), however, have species that occur in North Dakota. Members of the Pentatomidae can be recognized by the small to large size, the lack of spines on the tibiae, three tarsal segments, and the usually subtriangular scutellum (if scutellum is more spatulate, it still does not cover the entire abdominal dorsum).

Key to the Subfamilies of North Dakota Pentatomidae 1 Rostrum large, incrassate, first segment not fitting tidily between bucculae...... Asopinae – Rostrum smaller, slender, fitting neatly between bucculae...... 2 2012 THE GREAT LAKES ENTOMOLOGIST 321 2(1) Scutellum distinctly spatulate in shape; pronotum with distinct anterolaterally projecting tooth on each side just posterior to each compound eye; antehumeral tooth present; usually relatively flat, dark brown to gray in color...... Podopinae – Scutellum usually triangular in shape; if spatulate, then color usually paler, brown; pronotum lacking anterolaterad projecting tooth on each side near compound eyes, or each tooth very small; antehumeral tooth usually lacking...... Pentatominae

PENTATOMIDAE: ASOPINAE Spinola, 1850 The Asopinae include the predatory stink bugs; that is members of this subfamily have an enlarged rostrum, very little of which fits between the pair of bucculae on the ventral side of the head. Members of this subfamily feed on other insects, and several are considered to be important in biological control programs. There are at least nine species known (either from the literature or herein reported) from North Dakota, with another two possible.

Key to the Genera of North Dakota Asopinae 1 Apical part of scutellum (tongue) enlarged, as long as or longer than frenal margin of scutellum, and about as wide as corium...... Perillus – Apical part of scutellum small, shorter than frenal margin and much narrower than corium...... 2 2(1) Ostiole of scent gland without elevated ruga; peritreme terminating in short or long sulcus that is not surrounded by shagreened area of cuticle (evaporatorium)...... 3 – Ostiole of scent gland attended by elevated ruga surrounded by shagreened area of cuticle...... 4 3(2) Juga much longer than tylus, convergent, contiguous anteriorly [not yet found in North Dakota]...... Rhacognathus – Juga and tylus subequal in length [not yet found in North Dakota]...... Zicrona 4(2) Males lacking glandular patches of silky hairs on abdominal venter; female basal plates contiguous...... Podisus – Males with glandular patches of silky hairs on abdominal venter; female basal plates separated by intervening plate...... Apoecilus

Apoecilus Stål, 1870 Apoecilus was originally described as a subgenus of Podisus and is obvi- ously related to that genus. In fact, the species included here used to be in the genus Podisus, and then they were also included in another closely related genus Apateticus. The present arrangement was set forth by Thomas (1992). There are currently three species in Apoecilus, but A. invarius (Walker) occurs in the southwestern U.S. and Mexico. The remaining two species both occur in North Dakota, and can be separated by the following key.

Key to species of Apoecilus in North Dakota 1 Basal plates of female convergent apically, medial plate between them triangular; upper arm of male paramere curvilinear, not bent, and about equal in diameter to lower arm...... Apoecilus bracteatus (Fitch) 322 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

– Basal plates of female not convergent apically, medial plate between them quadrangular; upper arm of male paramere either bent or more slender in diameter than lower arm...... Apoecilus cynicus (Say) Apoecilus bracteatus (Fitch, 1856) - Interestingly, the two North Dakotan species of Apoecilus are easier to tell apart in the female sex than in males. The medial plate between the basal plates is triangular in this species, and squarish in A. cynicus. Males can be separated, however, by the male parameres. Each paramere is divided into a dorsal and a ventral arm. In this species, the two arms are nearly the same size and shape. In A. cynicus, the dorsal arm is often more curved and is more slender than the ventral arm. McPherson (1982) gave the distribution of this species (as Apateticus bracteatus) as Québec and New England west to Vancouver, Idaho, Colorado, New Mexico, Utah, and California. Although several published works have recorded this species from nearly all the states or provinces surrounding North Dakota (Van Duzee 1904, Froeschner 1988c, Maw et al. 2000), there is only a single published record from North Dakota. Hussey (1922) gave the following record: Bottineau Co.: Bottineau, T. H. Hubbell. The life history of this species was studied by Downes (1920, as Apateticus crocatus) and Evans and Root (1980, as Apateticus bracteatus). Specimens Examined: NORTH DAKOTA: Cass Co.: Fargo, 12-VII- 1967, J. W. Goertel (1♂ NDSU). Cavalier Co.: summer, 1958 (1♀ NDSU). Divide Co.: 26-VII-1958, S. Wilson (1♂ NDSU). Grand Forks Co.: Grand Forks, 4-VII-1970, G. Ruonala (1♀ UNDG), 28-VII-1966 (1♂ UNDG); Oakville Prairie, T151N R52W Sec. 16, 16-VII-1976 (2♀♀ UNDG); T151N, R50W, Sec. 16, 27-VI-1966 (1♀ UNDG). Hettinger Co.: 26-VIII-1976, Lago & Kurtz (1♂ NDSU). Stutsman Co.: VIII-1957 (1♀, NDSU). Apoecilus cynicus (Say, 1831) - This species is relatively easy to distinguish from A. bracteatus (see key and discussion above). McPherson (1982) gave the distribution for this species as Québec and New England south to Florida, and west to Montana, Colorado, Texas, and Arizona. Uhler (1878) listed two North Dakota records for this species: Bottineau Co.: near Mouse River, August 29, 1873, and Pembina Co.: plains near Pembina. The biology of this species (as Apateticus cynicus) was studied by Jones and Coppel (1963); they also described the immature stages. Specimens Examined: NORTH DAKOTA: Grand Forks Co.: Turtle R., 4.8 mi. W, 2 mi. N Emerado, T152N R53W Sec. 20 & 29, 19-IX-1984 (1♂ UNDG). Hettinger Co.: 12-VII-1976, Lago & Kurtz (1♂ NDSU). Richland Co.: Ekre Research Ranch, 48 32 15N, 27 12 00W, 9-VII-2002, A. Joshi (1♂ NDSU). Sargent Co.: 9mi. E, 1mi. S of Oakes, 23-IX-2007, K. Delahoyde (1♀ DARC).

Perillus Stål, 1862 This genus can be recognized by the somewhat enlarged scutellar tongue. The only other American genus with the scutellum enlarged is Stiretrus Laporte, but this genus does not occur in North Dakota. In Stiretrus, the scutellum is greatly enlarged, covering the majority of the abdomen. Besides the three species treated below, there are another four Perillus species not known from North Dakota. Perillus confluens Herrich-Schäffer is a southwestern U.S. species; P. lunatus Knight and P. splendidus (Uhler) occur in the western U.S.; and P. strigipes (Herrich-Schäffer) is a southeastern U.S. species. This genus was revised by Knight (1952) and Thomas (1992).

Key to Species of Perillus in North Dakota 1 Profemur with anteapical spine or stout tubercle; transverse pronotal dark band interrupted medially, forming two large spots...... 2 2012 THE GREAT LAKES ENTOMOLOGIST 323 – Profemur with tubercle obsolescent; transverse pronotal dark band entire, not interrupted medially...... Perillus exaptus (Say) 2(1) Abdominal spiracles enclosed within yellow area...... Perillus circumcinctus Stål – Abdominal spiracles enclosed within brown or black area...... Perillus bioculatus (Fabricius) Perillus bioculatus (Fabricius, 1775) - This species, commonly known as the twospotted stink bug, preys on a variety of species, but seems to have a preference for the larvae of Chrysomelidae, especially the Colorado potato beetle. It has been studied extensively as a biological control agent against that species. This species is known from two distinct color forms. The typical form is red and black; the less common form (given the name clanda Say) is white or yellowish with brown markings. McPherson (1982) gave the range of this species as Québec, Ontario, and New York west to the Pacific coast, and south to Florida, New Mexico, Arizona, and California. Uhler (1878), under the name Perillus claudus, recorded this species from near Turtle Mount [probably in Rollette Co.], July 24, 1873; later Hussey (1922) recorded P. bioculatus claudus from Devils Lake [Ramsey Co.], May, 1921, N. A. Wood. Knight (1923) studied the biology and life history of this species. Specimens Examined. NORTH DAKOTA: Burleigh Co.: Bismarck, IX-1921 (1♂ NDSU); VIII-1960 (1♀ NDSU). Cass Co.: Fargo, NDSU Campus, 46°54′42″N 96°48′14″W, 9-VII-2002, L. M. Tobias (1♀ NDSU); 28-VIII-1960 (1♂ NDSU). Divide Co.: Crosby, 5-IX-1969, T. D. Lindsey (1♀ UNDG). Grand Forks Co.: Grand Forks, 22-IX-1933 (1♀ UNDG), 22-IX-1941 (1♀ UNDG), IX- 1929 (1♂ UNDG), 1-X-1934, E. & G. Wheeler (1♀ UNDG), 3-X-1941 (1♂ UNDG), 4-X-1937 (1♂ UNDG), 28-X-1963 (1♂ DARC). McKenzie Co.: Watford City, 21-III-1985 (1♀ NDSU); 30-VIII-1978 (1♀ NDSU). McLean Co.: Garrison, Elevator, 10-VII-2007, K. Conklin (1♂ DARC); Washburn, 27-VII-1984, feeding on sunflower beetle larvae (1♂ NDSU); IX-1958 (1♂ 1♀ NDSU). Morton Co.: Mandan, 22-IX-1922, R. L. Webster (1♂ NDSU). Mountrail Co.: 13-VII-1957, P. Bergen (1♀ NDSU), 10-IX-1960 (1♂ NDSU); Stanley, 15-IX-1998, A. Nelson (1♀ MISU). Ward Co.: Minot, 10-IV-1936, H. Rosg (1♀ MISU), 5-V-1936, S. Anhalt (1♂ MISU), 9-V-1936 (1♀ MISU), 10-V-1936 (1♀ MISU), 13-VII-1937, T. Tate (1♀ MISU). Wells Co.: 6-IX-1959 (1♀, NDSU). Perillus circumcinctus Stål, 1862 - This species is very similar in appearance to the clanda form of P. bioculatus, but can be recognized by the lack of large dark spots surrounding the spiracles. McPherson (1982) gave the distribution of this species as Québec west to Saskatchewan, and south to New Jersey, Illinois, Missouri, and Nebraska. Uhler (1878) simply listed this species from “Dakota,” but Hussey (1922) provided two specific North Dakota collection records: Bottineau Co.: Bottineau, August 1, 1920, T. H. Hubbell, and Ramsey Co.: Devils Lake, July 28, 1920, T. H. Hubbell, on a raspberry bush feeding on a chrysomelid larva. Specimens Examined. NORTH DAKOTA: Burleigh Co.: 10-VII-61, A. C. Olson (1♀ NDSU). Cavalier Co.: Clyde, IX-1922, W. Baker (1? NDSU). Divide Co.: Crosby, 5-IX-1969, T. D. Lindsey (1♀ UNDG). Ransom Co.: 1-VIII- 1961 (1♂ NDSU). Perillus exaptus (Say, 1825) - This species is distinctly smaller than the previous two, and can be recognized by the continuous transverse pronotal dark band. Although Froeschner (1988c) listed this species from “Dakota,” I have not seen any specific literature records for North Dakota. McPherson (1982) gave the distribution as Québec and New England west to the Pacific coast, and south to New Jersey, Missouri, New Mexico, Utah, and California. Uhler (1878) provided a record from Montana, and Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. 324 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Specimens Examined. NORTH DAKOTA [New State Record]: McKen- zie Co.: Ted. Roos. Park, 24-VI-1964, R. J. Sauer (1♂ NDSU). Richland Co.: 2mi. NE Mirror Pool, 28-VI-1993, G. Fauske (1♂ NDSU); SNG, Horsetrail Head 46°31.409N, 97°12.194W, 25-VIII-2009, D. A. Rider (1♂ DARC). Slope Co.: Burning Coal Vein St. Pk., 10-VIII-1993, D. A. Rider (1♀ DARC).

Podisus Herrich-Schäffer, 1853 This is a large and diverse genus, especially in the Neotropics. Thomas (1992) has provided a review of the genus. There are six species known to occur in the United States, but one, P. sagitta (Fabricius), only enters the U.S. in southern Florida and Texas. Podisus neglectus seems to be an eastern U.S. species, although there are records as far west as Indiana and Michigan. The remaining four have been or are now recorded from North Dakota.

Key to Species of Podisus of North Dakota 1 Anterolateral pronotal margins straight; wing membrane clear (sometimes with a faint vitta)...... Podisus placidus Uhler – Anterolateral pronotal margins concave; wing membrane with a dark blotch or stripe...... 2 2(1) Humeri acute to spinose; femora with pair of anteapical spots; abdominal sternite VII with midventral spot...... Podisus maculiventris (Say) – Humeri rounded or obtusely angular; femora variably maculate or immaculate; abdominal sternites immaculate or with row of spots...... 3 3(2) Metafemora immaculate, except sometimes faint spot or two subapically; may or may not have a row of dark spots along midline of abdominal venter...... Podisus brevispinus Phillips – Metafemora densely spotted; usually with row of spots on midline of abdomen...... Podisus serieventris Uhler Podisus brevispinus Phillips, 1992 - This species has had a somewhat confused taxonomic history. Early workers treated this species under the name P. modestus (Dallas). Phillips (1983) in an unpublished dissertation determined that P. modestus was actually a synonym of P. maculiventris, thus leaving P. modestus of authors (not Dallas) without a name. She described it in her dissertation under the name P. brevispinus. Although Philipps never published her work in a reviewed journal, Thomas (1992) gave Phillips credit for her discoveries, and for the naming of this species. This species is most likely to be confused with P. maculiventris, but it can be recognized by the more rounded humeral angles, and the shorter medial abdominal spine. McPherson (1982, as P. modestus) gave the distribution of this species as Québec and New England west to British Columbia, Montana, and Idaho, and south to North Carolina, Georgia, and New Mexico. Although Froeschner (1988c, as P. modestus) listed this species from “Dakota,” Manitoba, and Montana, no specific localities from North Dakota have been published; Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. Phillips (1983) listed this species from Bot- tineau Co. and Towner Co. in her unpublished dissertation. The life history of this species (as Podisus modestus) was studied in Québec (Tostowaryk 1971), Wisconsin (Coppel and Jones 1963), and New York (Wheeler 1977). McPherson (1982, as P. modestus) provided a list of the nearly 50 prey species published for P. brevispinus. Specimens Examined. NORTH DAKOTA [New State Record]: Bottin- eau Co.: Turtle Mountains, 6-IV-1968, W. Kotchman (1♂ NDSU). Cavalier Co.: Langdon, 18-VIII-1929 (1♂ UNDG). Grand Forks Co.: Forest R. Biol. Area, 1.8mi. W, 1mi. N Inkster, T154N R55W Sec. 11 & 14, 18-IX-1976 (1♂ UNDG); 2012 THE GREAT LAKES ENTOMOLOGIST 325 Turtle River State Park, 3-VII-1965, S. Kraft (2♀♀ UNDG); University Biology Forest R. Valley Inkster, 18-VI-1958 (1♀ UNDG). Mercer Co.: T144N, R84W, Sec. 22, 4-6-VIII-1972, A. C. F. Hung (1♂ UNDG). Pembina Co.: Cavalier, 4-IX-1976 (1♀ UNDG). Richland Co.: Ekre Grassland Preserve, 46.552N, 97.130W, 28-V-2010, D. Rider (1♀ DARC), 3-IX-2009, D. A. Rider (4♂♂ 5♀♀ DARC); Mirror Pool, T135N, R52W, Sec 8, 29-VIII-1996, D. Rider (1♂ 3♀♀ NDSU); SNG, Horse trail head, 46°31.409N, 97°12.194W, 25-VIII-2009, D. A. Rider (1♀ DARC). Walsh Co.: S. Branch, Park River NE of Adams, 21-VIII- 1970, J. Singleton (1♀ UNDG). Podisus maculiventris (Say, 1831) - This species, commonly called the spined soldier bug, is relatively common throughout the U.S. It feeds on a variety of insect species. As mentioned above, it may be confused with P. brevispinus, but the humeral angles are always sharply spinose, and the ventral basal abdominal spine is longer, reaching between the hind coxae. It does not seem to be as common in North Dakota as P. brevispinus. Neither Uhler (1878) or Hussey (1922) (or anyone else) has previously recorded this species from North Dakota. McPherson (1982) gave the distribution as Québec and New England west to the Pacific coast, and south to Florida, Arizona, and California. Froeschner (1988c) listed this species from Manitoba and Montana, and Maw et al. (2000) recorded this species from Manitoba. Lugger (1900, as P. spinosus) recorded this species from Minnesota. Phillips (1983) listed this species from Grand Forks Co. in her unpublished dissertation. This species is generally considered to be a beneficial species in agricultural situations, even though it is an indescriminate predator (see list of prey species given by McPherson 1980, 1982). This has resulted in a multitude of published works on its biology and life history, including prey species, natural enemies, and biology in general (again, see McPherson 1982 for citations). Specimens Examined. NORTH DAKOTA [New State Record]: Billings Co.: Medora 47 02 00N, 103 36 00W, 24-VI-2002, A. Joshi (1♂ NDSU). Cass Co.: Fargo, NDSU campus, 5-VIII-2002, D. Doyle (1♂ NDSU), 5-IX-2000, R. Aylesworth (1♂ NDSU); SW Fargo, 1-X-2002, S. Koudela (1♂ NDSU); Wild Rice R. at Red Riv., 46.758N, 96.787W, 1-VIII-2012, D. Rider (1♂ DARC), 13-VIII- 2012, D. A. Rider (1♀ DARC); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (2♀♀ DARC), 1-VIII-2012, D. A. Rider (4♀♀ DARC), 13-VIII-2012, D. A. Rider (1♂ DARC). Grand Forks Co.: Northwood, 11-VII- 1953, R. L. Post (1♀ NDSU); Oakville Prairie, T151N R53W Sec. 16, 14-IX-1977 (1♂ UNDG). Ransom Co.: SNG, Mirror Pool, T135N R53W SW Sec. 1, 22-VII- 2002, L. Tobias (2♂♂ NDSU). Richland Co.: Mirror Pool, 26-VIII-1994, T. Nowatzki (1♂ NDSU); SNG, Oak Savannah Area, 18-VI-2000, D. A. Rider (1♀ DARC). Ward Co.: Minot, 10-VII-2002, G. Hanley (1♀ MISU). Podisus placidus Uhler, 1870 - This is the easiest species of Podisus to identify in North Dakota. The straight anterolateral pronotal margins, and the usually clear membrane, will separate this species from the other Podisus species. McPherson (1982) gave the distribution as Québec west to British Columbia, and south to New Jersey, Arkansas, Colorado, Utah, and Idaho. Froeschner (1988c) did not list this species from North Dakota, or from any of the surrounding states or provinces (Alberta, Ontario, Iowa are some of the closest records), and I have not seen any other published records from either North Dakota or Minnesota. Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. Phillips (1983) listed this species from both Minnesota and North Dakota in her unpublished dissertation: MINNESOTA: Carlton Co., Cass Co., Clearwater Co., Hennepin Co., Mille Lacs Co., Olmstead Co., Ottertail Co., Pine Co., Ramsey Co., St. Louis Co., Wabash Co. and the three NORTH DAKOTA records are included in the data below (she examined the NDSU specimens from Bill- ings Co., Cass Co.: Fargo, Golden Valley Co.). The life history and immature stages have been described (Coppel and Jones 1963, Oetting and Yonke 1971a). McPherson (1982) provided a list of known prey species. 326 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Specimens Examined. MINNESOTA [New State Record]: Cleawater Co.: Itasca Pk., 17-VI-1937, H. R. Dodge (1♂ DARC), 8-VIII-1937, H. R. Dodge (1♀ DARC). St. Louis Co.: Dark Lake Barrens, T60N R20W 25, 31-V-1992, T. Warner (1♀ NDSU). NORTH DAKOTA [New State Record]: Billings Co.: Theo. Roos. Mem. Pk., 30-VIII-1961, R. L. Post (1♀ NDSU). Cass Co.: Fargo, 14-VI-1923, R. L. Webster (1♀ NDSU); Leonard, 22-VII-1989, D. Hyder, on sunflower (1♀ NDSU). Golden Valley Co.: Beach, 19-IX-1922, R. L. Webster (1♂ NDSU). Richland Co.: 1mi. E Mirror Pool, 22-VII-1993, D. A. Rider, reared from nymph (2♂♂ DARC). Podisus serieventris Uhler, 1871 - It may be difficult to separate this species from P. brevispinus. In well marked specimens, it should be no trouble. The hind femora in P. serieventris should have numerous dark spots, and there should be a row of dark spots along the midline of the abdomen. I have seen specimens of P. brevispinus, however, that also have the black spots on the abdominal venter, but they do not have the hind femora maculate as in this species. McPherson (1982) gave the range for this species as Nova Scotia and Québec west to British Columbia, and south to North Carolina, Florida, New Mexico, Utah, and Idaho. Froeschner (1988c) listed this species from Minnesota and Montana, and Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. Phillips (1983), in her unpublished dissertation, reported examining the specimen from Cass Co. that is listed below. The biology of this species has been studied (Prebble 1933), and some limited information about this species life history was given by Coppel and Jones (1963). Specimen Examined. NORTH DAKOTA [New State Record]: Cass Co.: Fargo, VI-1950, G. J. & R. L. Post (1♀ NDSU).

Rhacognathus Fieber, 1860 Rhacognathus is generally considered to be an Old World genus with a single species known from North America. Rhacognathus americanus Stål occurs in southeastern Canada and northeastern United States. Van Duzee (1904) recorded this species from Winnipeg, Manitoba; Froeschner (1988c) listed this species from Manitoba and Minnesota; and Maw et al. (2000) recorded this species from Manitoba. It is likely that this species will eventually be collected in North Dakota, probably in eastern or northern areas of the state.

Zicrona Amyot and Serville, 1843 This genus contains two North American species, the cosmopolitan Z. caerulea (L.) and the recently described Z. americana Thomas. The latter species occurs in the southwestern and western states. Froeschner (1988c) did not list Z. caerulea from North Dakota, or from any of the surrounding states or provinces; but Maw (2000) recorded this species from Manitoba and Saskatchewan. It may eventually be collected in North Dakota.

PENTATOMIDAE: PENTATOMINAE Leach, 1815 The Pentatominae is the largest subfamily of stink bugs, and all known species are phytophagous. This subfamily contains many of the economically important pests. I have provided two keys for the reader to use to identify their specimens to the generic level. First, I have provided a key to the tribes known to occur in North Dakota. Once you have finished with this key, you will then find a key to the genera of each tribe located under each tribal heading. Second, following the key to tribes, I have also provided a key to all pentatomine genera known to 2012 THE GREAT LAKES ENTOMOLOGIST 327 occur in North Dakota, regardless of what tribe they occur in. The two keys are similar in some ways, but different in others; the hope being, one or the other will get the reader to the correct identity.

Key to the Tribes of North Dakota Pentatominae 1 Lateral margin of each jugum with a relatively conspicuous tooth or denticle between the compound eye and the apex, usually near apex; anterolateral pronotal margins coarsely denticulate; veins in wing membrane dark brown with brown and white pattern between veins...... Halyini – Apex of each jugum may be acute or toothlike, but lateral margin lacking conspicuous tooth between compound eye and apex; anterolateral pronotal margins variable, but never coarsely denticulate; veins in wing membrane variable, but never with bicolored pattern between veins...... 2 2(1) Coloration pale yellow; elongate, slender, length about four times width; abdominal venter with stridulatory area on first three sternites on each side of midline...... Mecideini – Coloration variable, but if pale yellow, then more broadly rounded (some fall forms of Thyanta custator accerra) or humeral angles spinose (Oebalus pugnax, which is not known to occur in North Dakota); not especially elongate or slender, no more than twice as long as width; stridulatory areas on abdominal venter lacking...... 3 3(2) Ostiole very small, difficult to see, located mesially between middle and hind coxae, and lacking attending ruga, canal, or evaporative area...... Strachiini – Ostiole larger, more conspicuous, located laterad of coxal bases, and always attended by at least a small evaporative area, and usually a canal or ruga of various sizes and/or shapes...... 4 4(3) Relatively small, flattened, with lateral margins of head, pronotum, and coria explanate...... Sciocorini – Usually larger, if small, then lateral margins of head, pronotum, and coria not explanate...... 5 5(4) Anterior margin of propleura explanate, convex, expanded anteroventrad up to the base of the antennifer, sometimes covering the antennifer and the base of the first antennal segment...... Aeliini – Anterior margin of propleura not explanate, usually more or less straight, remote from the antennifer...... 6 6(5) Primarily green in color, at most with a few small black or yellow spots or stripes, and remaining green after death...... Nezarini – Color variable, but usually not mostly green...... 7 7(6) Base of abdominal venter produced anterad as a small spine or tubercle...... Pentatomini – Base of abdominal venter not produced anterad, smoothly rounded...... Carpocorini Key to the Genera of North Dakota Pentatominae 1 Body strongly pubescent, dorsal surface with many long, bristle-like hairs [Carpocorini (part)]...... Trichopepla – Body not strongly pubescent, perhaps a few longer hairs on legs, but dorsal surface nearly glabrous (Brochymena and Parabrochymena may have small, white, scale-like hairs, but never long bristle-like hairs)...... 2 328 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

2(1) Elongate, slender, length about four times width, coloration pale yellow; abdominal venter with stridulatory area on first three sternites on each side of middle [Mecideini]...... Mecidea – Not particularly elongate or slender, no more than twice as long as width, coloration variable, but if yellowish, then usually more broadly rounded (some specimens of the fall form of Thyanta) or humeral angles are spinose (Oebalus pugnax which is not yet known from North Da- kota); stridulatory areas on abdominal venter lacking...... 3 3(2) Predominant color black with red or orange markings, or orange with black markings (rarely orange coloration faded in preserved specimens and appear pale yellow to white)...... 4 – Predominant color brown, gray, or green; if nearly black, then lacking red or orange markings...... 5 4(3) Relatively small, length less than 8mm, outline nearly circular; dorsal coloration black with red cross-shaped marking on pronotum and a pair of red spots near apex of scutellum; ostiole and accompanying ruga or canal conspicuous [Carpocorini (part)]...... Cosmopepla – Medium in size, length 8 to 12mm, shape somewhat more elongate-oval, dorsal color usually orange with black markings, occasionally more black than orange; ostiole and accompanying ruga or canal obsolete [Strachiini]...... Murgantia 5(3) Lateral margin of each juga with a conspicuous tooth or denticle between apex and compound eye, usually near apex; anterolateral pronotal margins coarsely denticulate; veins in hemelytral membrane dark brown with a brown and white pattern in the membrane between veins [Halyini]...... 6 – Each juga may be acute apically, but lateral margin never with a tooth or denticle between apex and compound eye; anterolateral pronotal margin variable, but never coarsely toothed; veins in hemelytral membrane variable, but never with brown and white pattern in membrane between veins...... 7 6(5) Humeral angles subquadrate; basal disk of scutellum distinctly elevated...... Parabrochymena – Humeral angles subtriangular, rounded to angulate, but not quadrate; basal disk of scutellum at most slightly elevated...... Brochymena 7(5) Dorsal coloration predominantly green, sometimes with yellowish stripes or spots, and/or small black spots...... 8 – Dorsal coloration predominantly brown or black, often with pale markings...... 12 8(7) Base of abdominal venter produced forward into a small obtuse spine or tubercle...... 9 – Base of abdominal venter smoothly rounded, not produced forward [Nezarini (part)]...... 10 9(8) Smaller, less than 15mm in length [Pentatomini (part)]...... Banasa (part) – Larger, more than 25mm in length [Nezarini (part)]...... Chinavia 10(8) Smaller, usually less than 8mm in length...... Tepa – Larger, usually more than 10mm in length...... 11 11(10) Ostiolar ruga elongate, reaching at least two-thirds to edge of supporting plate, and acuminate apically...... Thyanta 2012 THE GREAT LAKES ENTOMOLOGIST 329 – Ostiolar ruga much shorter, reaching no more than half way to edge of supporting plate, not acuminate apically...... Chlorochroa 12(7) Relatively small, body broadly oval , somewhat flattened, lateral margins of pronotum and corium explanate; width of body greatest across abdomen; head nearly as broad as basal width of scutellum [Sciocorini]...... Sciocoris – Larger, if small, then body not unusually broad, usually more robust, lateral margins rounded, carinate, or reflexed, but not explanate; width of body usually greatest across pronotum, sometimes across abdomen; head usually not as wide as basal width of scutellum...... 13 13(12) Base of abdominal venter with small basal spine or tubercle basally [Pentatomini (part)]...... Banasa (part) – Base of abdominal venter smoothly rounded basally...... 14 14(13) Ostiolar ruga relatively short, more or less auriculate (maybe lacking or very small in Neottiglossa)...... 15 – Ostiolar ruga relatively long, not auriculate, tapering laterally, forming acute to acuminate apex...... 21 15(14) Scutellum subtriangular, shorter than corium, with apical third narrower than apex of corium, apex narrowly rounded [Carpocorini (part)]...... 16 – Scutellum more spatulate, longer than corium, with apical third wider than apex of corium, apex broadly rounded...... 19 16(15) Relatively small, less than 7.5mm in length; dark chocolate brown with some ivory markings on scutellum and a transverse ivory callus on anterior pronotal disk...... Mormidea – Larger, usually more than 8mm in length; color variable, but never chocolate brown, and never with transverse ivory callus on anterior pronotal disk...... 17 17(16) Anterolateral pronotal margins crenulate or dentate; humeral angles often prominent, angulate to spinose (rounded in Euschistus tristigmus)...... Euschistus – Anterolateral pronotal margins entire, not crenulate or dentate; humeral angles broadly rounded...... 18 18(17) Anterolateral pronotal margins somewhat explanate, distinctly convex; overall color grayish...... Menecles – Anterolateral pronotal margins not explanate, straight to slightly convex; overall color yellowish-brown, sometimes with reddish hues...... Antheminia 19(15) Head horizontal, or slightly declivent, tylus distinctly elevated above juga, juga not meeting anterior to tylus; propleura not produced anteroventrad, remote from adjacent antennifer [Carpocorini (part)]...... Coenus – Head strongly declivent, tylus at most slightly elevated above juga, juga longer than and contiguous anterior to tylus; propleura explanate, convex, produced anteroventrad, contiguous with or covering antennifer, and sometimes first antennal segment [Aeliini]...... 20 20(19) Relatively small, less than 7mm in length; prostethium with anterior margin not extending beyond anterior margin of eye, contiguous with or covering base of antennifer...... Neottiglossa – Larger, usually greater than 8mm in length; prostethium with anterior margin extending beyond anterior margin of eye, covering antennifer and at least part of first antennal segment...... Aelia 330 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

21(14) Juga distinctly longer than tylus and often contiguous anterior to tylus...... Holcostethus – Juga and tylus subequal in length or juga slightly longer, never contiguous anterior to tylus...... Halyomorpha

Tribe Aeliini Douglas and Scott, 1865 Members of this tribe are characterized by the produced prostethium which covers or nearly covers the base of the antennifers; they also tend to be somewhat cylindrical in shape. This is a relatively small tribe, at present containing only three genera. Aelia and Neottiglossa occur in both the New World and Old World, but both are limited to the Nearctic and Palearctic regions. The third genus, Aeliopsis occurs in northern Africa. Members of both Aelia and Neottiglossa are primarily grass feeders.

Key to the genera of Aeliini of North Dakota 1 Relatively small, less than 7mm in length; prostethium with anterior margin not extending beyond anterior margin of eye, contiguous with or covering base of antennifer...... Neottiglossa – Larger, usually greater than 8mm in length; prostethium with anterior margin extending beyond anterior margin of eye, covering antennifer and at least part of first antennal segment...... Aelia

Aelia Fabricius, 1803 Aelia is primarily an Old World genus with a single species occurring in North America. Several species are known to cause serious damage to wheat in Europe and the Middle East, but our species seems to prefer non-agricultural prairie grasses. It has been reported occasionally from wheat, but damage has been minimal. Aelia americana Dallas, 1851 - This species is fairly distinctive. It is cylindrical in shape, a little larger than the species of Neottiglossa, and the prostethium is greatly expanded to cover the adjacent antennifer. The range of this species is given by McPherson (1982) as British Columbia south to Arizona, and east to Manitoba, Michigan, and Illinois. Rider (1986a) extended the known range southeastward to Arkansas and Alabama. Although Froeschner (1988c) and Maw et al. (2000) recorded this species from all states or provinces north, west, and south of North Dakota, it appears that no one has recorded it from North Dakota or Minnesota. Froeschner (1942) studied the immature states of this species, and compared them with those of Neottiglossa species. Specimens Examined. MINNESOTA [New State Record]: Clay Co.: Blazing Star Prairie, 47° 03′Lat 96° 25′Long., P. Tinerella & G. Fauske, pitfall (1♀ NDSU); Bluestem Prairie, 46° 50′Lat. 96° 26′ Long., 5-VIII-1996, C. Locken & G. Fauske, pitfall (1♂ 2♀♀ NDSU), 3-IX-1996, C. Locken & T. Kochis, pitfall (1♀ NDSU); 46° 51′Lat. 96° 28′Long., 27-VII-1995, D. Rider & G. Fauske, pitfall (1♀ NDSU), 31-VII-1997, P. Tinerella & A. Abbott, pitfall (1♀ NDSU), 26-VIII- 1995, G. Fauske & D. Rider, pitfall (1♀ NDSU), 16-IX-1996, C. Locken & T. Kochis, pitfall (1♀ NDSU); Bluestem SNA, T139N R46W Sec. 23, 27-VI-1996, C. Locken & G. Fauske, pitfall (1♀ NDSU); CCL Trust Lands, 47° 03′Lat. 96° 26′Long., 24-VI-1997, P. Tinerella & G. Fauske, pitfall (2♀♀ NDSU), 3-VII-1996, J. Albertson & L. DeCock, pitfall (1♀ NDSU), 9-VII-1997, P. Tinerella, pitfall (1♂ NDSU), 13-VII-1999, P. Tinerella & G. Fauske, pitfall (1♀ NDSU), 1-VIII- 1996, C. Locken & K. Urlacher, pitfall (1♀ NDSU), 28-VIII-1997, P. Tinerella & A. Abbott, pitfall (1♀ NDSU). Polk Co.: Agassiz Dunes SNA, 47° 30′Lat. 96° 18′Long., 21-VIII-1997, G. Fauske & P. Tinerella, pitfall (1♀ NDSU). NORTH 2012 THE GREAT LAKES ENTOMOLOGIST 331

DAKOTA [New State Record]: Barnes Co.: Natl. Fish Hatchery, 7-VI-1978, I. Nashnosh (1♂ NDSU); Valley City S.U., 17-X-1976 (1♀ UNDG). Billings Co.: Medora, T141N R102W, 24-VI-2002, L. M. Tobias (1♀ NDSU). Cass Co.: 4-X-1954, B. Bernardy (1♀ NDSU). Emmons Co.: Linton, 22-VII-1964, R. J. Sauer (1♂ NDSU). Grand Forks Co.: Arvilla, J. M. Davis (1♂ UNDG); Oakville Prairie, Sec. 9 Oakville Township, 5-VII-1961 (1♂ UNDG), 3-X-1959 (1♂ UNDG). McKenzie Co.: T.R. Nat. Pk., NU, 23-VI-1977, E. U. Balsbaugh & A. Jamieson (1♀ NDSU). Mercer Co.: Sec. 22, T144N, R84W, 4-6-VIII-1972, A. C. F. Hung (1♀ UNDG). Richland Co.: SNG, 2.5mi. SE McLeod, 26-V-1992, D. Rider (1♂ DARC). Ward Co.: NW Minot, 6-VI-1994, G. Hanley (1♀ MISU); SE Minot, 17-V-1994, G. Hanley (1♀ MISU), 22-V-1994, G. Hanley (1♀ MISU). Williams Co.: Williston, 23-VIII-1987, M. J. Weiss (2♀♀ NDSU).

Neottiglossa Kirby, 1837 This genus has species occurring in both North America and Europe and Asia. The North American species were recently revised (Rider 1990). They are relatively small, somewhat cylindrical species, usually brown to fuscus with some pale markings; they can be separated from Aelia by their smaller size, and that the expanded prostethium at most covers only the base of the antennifer. Only two species are known from North Dakota, but a third, N. trilineata (Kirby), may eventually be found in the state. It has been recorded from Manitoba, Saskatchewan, and “Dakota” (Froeschner 1988c, Maw et al. 2000). The three species can be separated by the following key.

Key to species of Neottiglossa that may occur in North Dakota 1 Coxae pale yellow; evaporative surfaces pale yellow to brown-gray with contrasting black punctures...... 2 – Coxae fuscous to black; evaporative surfaces black with concolorous punctures...... Neottiglossa sulcifrons Stål 2(1) Dorsal surface of head and propleura mostly black with concolorous punctures [not yet recorded from North Dakota]...... Neottiglossa trilineata (Kirby) – Dorsal surface of head and propleura with large areas of pale yellow to brown with black punctures...... Neottiglossa undata (Say) Neottiglossa (Texas) sulcifrons Stål, 1872 - This species can be separated from the other two species that may occur in North Dakota by the dark coxae and evaporative surfaces. Blatchley (1926) gave the range of this species as New Jersey south to Georgia, and west to Nebraska, Kansas, and New Mexico; McPherson (1982) indicated that the known range should be extended to South Dakota and British Columbia. This species has not been reported from North Dakota; in fact, neither Froeschner (1988c) nor Maw et al. (2000) listed this species from any of the surrounding states or provinces. The immature stages have been described (Esselbaugh 1946, DeCoursey and Esselbaugh 1962). Specimens Examined: MINNESOTA [New State Record]: Clay Co.: 3mi. E, 2mi. S Felton, 47°03.71′N, 96°25.99′W, 26-VI-2002, P. B. Beauzay (1♂ NDSU). NORTH DAKOTA [New State Record]: Billings Co.: Freeman’s Fron- tier near Medora, 9-VIII-1993, D. A. Rider (1♀ DARC); Medora, 47°02′00″N, 103°36′00″W, 24-VI-2002, A. Joshi (2♂♂ NDSU). Oliver Co.: T43N, R83W, Sec. 6, 15-VIII-1972, P. B. Kannowski (1♂ 2♀♀ UNDG). Richland Co.: Mirror Pool, T135N, R52W, Sec. 8, 27-VI-1992, D. A. Rider (1♂ DARC). Neottiglossa (Neottiglossa) undata (Say, 1832) - This species tends to be slightly larger than N. sulcifrons, and it has the coxae pale, as well as the 332 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 evaporative area (with contrasting dark punctures). McPherson (1982) gave the known distribution as Québec and New England west across southern Canada and the northern United States to the Pacific coast. There is a single published record of this species from North Dakota. Hussey (1922) recorded this from near the Bois de Sioux River at Fargo (Cass Co.). Froeschner (1988c) listed this species from Minnesota, and Maw et al. (2000) recorded it from both Manitoba and Saskatchewan. Specimens Examined: NORTH DAKOTA: Bottineau Co.: 4-VI-1968, R. W. Katayama (1♂ NDSU). Cass Co.: Fargo, 31-VIII-2004, S. Kuchera (1♀ NDSU); S Fargo, 76 Ave. S, Chrisan Edition, 46.766N, 96.780W, 5-VI-2012, D. A. Rider (1♂ DARC); Wildlife Man. Area N of Brewers Lake, 26-V-2010, D. A. Rider (1♀ DARC); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (1♂ 1♀ DARC), 1-VIII-2012, D. A. Rider (2♀♀ DARC). Dickey Co.: 22-VI-1959, R. L. Post (1♀ NDSU). Grand Forks Co.: Air Force Base Prop- erty, McKinock Twp, T152N, R53W, Sec. 27, 16-IX-1970 (1♂ UNDG); Arvilla, 30-IX-1929 (1♂ UNDG); Grand Forks, 21-VII-1958 (1♀ UNDG); Northwood, 15-VI-1964, R. L. Post (1♀ NDSU); Oakville Prairie, Oakville Twp, Sec. 9, 28- VI-1961 (1♂ UNDG); Turtle River, 5mi. W, 2mi. N Emerado, 8-IX-1975 (1♀ UNDG); Turtle River State Park, 30-VI-1953 (1♂ UNDG); UNDG Biology Area, Forest R. Valley, near Inkster, 12-VII-1963 (1♀ UNDG), 27-IX-1968, R. S. Tro- bec (1♀ UNDG). McKenzie Co.: Th. Roos. M. Prk, 22-VI-1965, W. Kotchman (1♂ NDSU). Mercer Co.: T144N, R84W, Sec. 22, 4-6-VIII-1972, A. C. F. Hung (1♀ UNDG). Oliver Co.: T143N, R83W, Sec. 6, 15-VIII-1972, P. B. Kannowski (2♀♀ UNDG). Ransom Co.: Mirror Pool WMA, 46°32.348N, 97°17.475W, 27- VI-2012, D. A. Rider (2♀♀ DARC); SNG, A-Annex, T136N, R52W, Sec. 8, 4-IX- 1992, D. A. Rider (1♀ DARC); SNG, 1.5mi. W McLeod, T134N, R53W, Sec. 22, 4-VI-1992, D. A. Rider (1♂ DARC). Richland Co.: Ekre Grassland Preserve, 22-VII-1993, D. L. Cuthrell (1♀ NDSU), 25-VII-1995, G. Fauske (1♂ NDSU), 28-VIII-2008, D. A. Rider (1♀ DARC), 3-IX-2009, D. A. Rider (1♂ DARC), 25- IX-1994, T. Nowatzki (1♂ 1♀ NDSU); Mirror Pool, T135N, R52W, Sec. 8, 27- VI-1992, D. A. Rider (1♂ 1♀ DARC); 6-VI-1962, R. D. Frye, alfalfa (1♂ NDSU); SNG, Oak Savannah, 17-VII-1993, D. Cuthrell (2♀♀ NDSU); SNG, Horse trail head, 46.52N, 96.21W, 20-VII-2012, D. A. Rider (2♀♀ DARC); SNG, Horsetrail Head, 46°31.409N, 97°12.194W, 25-VIII-2009, D. A. Rider (1♂ 2♀♀ DARC). Ward Co.: Minot, Roosevelt Pk., 15-V-1994, G. Hanley (1♀ MISU); SE Minot, 23-V-1994, G. Hanley (1♀ MISU), 28-V-1994, G. Hanley (1♀ MISU). Williams Co.: D. A. Mundal, Euphorbia esula (1♂ NDSU).

Tribe Cappaeini Atkinson, 1888 At present, this tribe contains 23 genera, none of which is native to the New World. The majority of included genera occurs in Africa, but several genera can also be found in the Indian and Oriental regions, and even out into the Austra- lian region. This tribe is included here simply because of a recent introduction of one of its members into the New World. Halyomorpha halys (Stål) was first discovered around 1998 in eastern Pennsylvania (Hoebeke and Carter 2003). Its range has spread extensively in the eastern U.S. and a number of disjunct populations have now been discovered. No known populations exist in North Dakota, but individual specimens have been found (in shipping containers) in Grand Forks (Grand Forks Co.) and from western Minnesota (Clay Co.). It may be only a matter of time before this nuisance species arrives and becomes established here.

Tribe Carpocorini Mulsant and Rey, 1866 In general, this tribe contains those pentatomine genera which lack a tubercle or spine on the abdominal venter, and have auriculate ostiolar rugae. 2012 THE GREAT LAKES ENTOMOLOGIST 333 Occasionally, the ostiolar rugae may be elongate, but rarely are they acuminate as seen in the Antestiini or some of the Nezarini or Pentatomini. Many of our genera have traditionally been placed in the Pentatomini. This is the largest pentatomine tribe, containing many genera; it is also the most cosmopolitan, occurring in all geographical regions.

Key to Genera of Carpocorini known to occur in North Dakota 1 Dorsal surface of body covered with elongate, erect hairs.Trichopepla – Dorsal surface of body nearly glabrous...... 2 2(1) Overall coloration black with reddish markings forming a cross on the pronotum and two small spots near apex of scutellum...... Cosmopepla – Not colored as above, usually pale yellow to brown, and usually lacking reddish markings...... 3 3(2) Small, dark brown with an transverse ivory line (callus) on pronotum...... Mormidea – Usually larger, not colored as above...... 4 4(3) Scutellum somewhat spatulate in shape, apex broadly rounded...... Coenus – Scutellum subtriangular in shape, apex narrowly rounded...... 5 5(4) Ostiolar rugae elongate, tapering to a subacute apex; juga longer than and usually meeting in front of tylus...... Holcostethus – Ostiolar rugae auriculate, not subacute apically; juga and tylus usually subequal in length, but if juga longer (Euschistus servus), usually separated apically...... 6 6(5) Anterolateral pronotal margins crenulate or dentate; humeral angles usually prominent, angulate to spinose (rounded in Euschistus tristigmus)...... Euschistus – Anterolateral pronotal margins entire, not crenulate or dentate; humeral angles usually broadly rounded...... 7 7(6) Anterolateral pronotal margins somewhat explanate, distinctly convex; overall color grayish...... Menecles – Anterolateral pronotal margins not explanate, straight to slightly convex; overall color yellowish-brown, sometimes with reddish hues...... Antheminia

Antheminia Mulsant and Rey, 1866 Antheminia is mainly an Old World genus with two species known from North America. Our two species were treated by Thomas (1974) under the generic name Codophila. There has been some discussion, however, that the Pacific coast species, A. sulcata (Van Duzee), may only be a subspecies of A. eurynota. The single species in our area is also a western species, just reaching into the western half of North Dakota. The nominate subspecies occurs in the Old World in eastern Russia. Antheminia eurynota remota (Horváth, 1908) - The taxon remota was considered to be a valid species until Kerzhner (1993) determined that it was conspecific with the Old WorldA. eurynota. He continued to recognize the North American form as a subspecies of A. eurynota. This is a western U.S. species, occurring only in the western half of North Dakota. This species was not treated by McPherson (1982). Hussey (1922, as Carpocoris remota) provided the first two records of this species from North Dakota: Slope Co.: prairie near Amidon, 334 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

23-VIII-1920, T. H. Hubbell, and Williams Co.: Buford, A. L. Olson. Van Duzee (1904, as Carpocoris lynx) recorded this species from Williams Co.: Williston. Specimens Examined. NORTH DAKOTA: Billings Co.: Easy Hill, T138N, R100W, D. Aarhus (1♂ NDSU); Freeman’s Frontier near Medora, 9-VIII-1993, D. A. Rider, on Chenopodiaceae (1♀ DARC); Historical Marker, S. Unit, T. 140, R. 02, S. 10, Port. 400, 22-VI-1977 (1♀ NDSU). Eddy Co.: 22- VIII-1958, S. Wilson (2♀♀ NDSU). Golden Valley Co.: 25-VI-1957, P. Bergen (1♀ NDSU). McLean Co.: Ruso, 3-VII-1938, E. Boe (1♂ MISU). Mercer Co.: Sec. 9, T144N, R85W, 18-20-VII-1972, A, C. F. Hung (1♂ UNDG). Mountrail Co.: 12-VII-1957, P. Bergen (1♀ NDSU). Slope Co.: Burning Coal Vein St. Pk., 12-VIII-1993, D. A. Rider (1♂ DARC). Ward Co.: Minot, 10-V-1936, Hovdestad (1♀ MISU). Williams Co.: Williston, 28-VI-1987, M. J. Weiss (2♀♀ NDSU); Williston Exp. Stat., 1mi. W of Williston, IV-IX-1986, M. J. Weiss, pitfall (1♂ 1♀ NDSU); 11-VII-1961, V. Goodfellow, wheat (1♀ NDSU).

Coenus Dallas, 1851 Members of the genus Coenus tend to be more ovate, or even somewhat tear-drop in shape; the scutellum is enlarged, more spatulate than in most pentatomids, but not as enlarged as in the scutellerids or thyreocorids. There are currently three species in the genus. Coenus inermis Harris and Johnston occurs in the south central states with the closest records to North Dakota being from Missouri; C. explanatus Rider was recently described from southern Alabama and Georgia. The genus was recently reviewed (Rider 1996). Coenus delius (Say, 1832) - This species is relatively easy to recognize by the robust teardrop shape and the broadly rounded scutellum. McPherson (1982) gave the known distribution as Québec and New England south to North Carolina, and west to British Columbia, Montana, Idaho, Utah, Oklahoma, and Texas. This species probably occurs statewide in North Dakota. Uhler (1878) reported collecting it in two localities: Bottineau Co.: Mouse River region, August 29, and Pembina Co.: vicinity of Pembina. Hussey (1922) added several more records: Bottineau Co.: Turtle Mountains, T. H. Hubbell, Eddy Co.: Sheyenne River, about 3 miles S of Warwick, August 8, 1920, A. L. Olson, Nelson Co.: Stump Lake, T. H. Hubbell, Ramsey Co.: Devils Lake, T. H. Hubbell, and Slope Co.: Amidon, T. H. Hubbell. Rider (1996) presented a number of additional records from North Dakota, including abbreviated information (counties and a few specific localities); the full data is presented below. The life history of this species has been studied (Oetting and Yonke 1971b), and the immature stages have been described (Esselbaugh 1946, DeCoursey and Esselbaugh 1962). Specimens Examined: Benson Co.: 9-VIII-1958, S. Wilson (1♀ NDSU). Billings Co.: Easy Hill, T138N, R100W, D. Aarhus (1♂ 1♀ NDSU); Fremman’s Frontier near Medora, 9-VIII-1993, D. A. Rider (1♀ DARC); Medora, T141N, R102W, 24-VI-2002, L. M. Tobias (1♀ DARC); Th. Roos. M. Prk, 29-VI-1965, L. Grochowski (2♀♀ NDSU), 15-VII-1965, W. Kotchman (1♀ NDSU), 11-VIII- 1965, W. Kotchman (1♂ NDSU); 3-VII-1966 (2♀♀ NDSU). Bottineau Co.: Omemee, 29-VII-2008, D. Larson (1♀ NDSU). Cass Co.: Fargo, 1-V-1939, E. Sulerud (1♀ NDSU), 30-V-1940 (1♀ NDSU), VII-1966, J. W. Goertel (1♂ NDSU); Fargo, NDSU campus, 14-IX-2002, N. Boll (1♂ NDSU); 20-IX-1958, Roeshans (1♂ NDSU), S. Wilson (2♀♀ NDSU). Dunn Co.: 146-97-25-400, 13-VII-1976, Lago & Kurtz (1♀ NDSU), 20-VII-1977, L. A. Schutz & J. W. Smith (1♂ NDSU). Grand Forks Co.: Air force Base Property, McKinock Twp, Sec. 27, T152N, R53W, 18-IX-1974 (1♂ UNDG); Forest River Biology area, T154N, R55W, S11&14, 2-VII-1953 (1♀ UNDG); Grand Forks, 9-VII-1953 (1♀ UNDG), 21-IX- 1931 (1♀ UNDG), 13-X-1947 (1♀ UNDG); Kelly’s Slough NWR, 18-VI-1993, D. L. Cuthrell (1♀ DARC); Oakville Prairie, Sec. 9, T151N, R52W, Oakville Township, 22-VI-2012, D. A. Rider (3♂♂ 3♀♀ DARC), 23-VI-1961 (1♀ UNDG), 26-VI-1961 2012 THE GREAT LAKES ENTOMOLOGIST 335 (1♀ UNDG), 28-VI-1961 (2♀♀ UNDG), 5-VII-1961 (1♀ UNDG), 5-VII-1962, C. Sweikert (1♀ UNDG), 31-VII-1961 (1♂ 1♀ UNDG), 15-IX-1975 (2♀♀ UNDG), 5-X-1965 (1♀ UNDG); Oakville Prairie, Sec. 16, 47°53.403N, 97°18.855W, 22- VI-2012, D. A. Rider (1♂ 8♀♀ DARC), 21-VIII-2009, D. A. Rider (2♀♀ DARC), 24-VIII-2009, D. A. Rider (1♀ DARC), 26-IX-1983, K. Howe (2♂♂ UNDG), 4-X- 1965 (1♀ UNDG); Turtle R, 4.8mi. W, 2mi. N Emerado, T152N, R53W, Sec. 20&29, 8-IX-1982, K. Kreil (1♂ UNDG), 18-IX-1976 (1♂ UNDG); T151N, R50W, 1-X-1965 (1♀ UNDG). McHenry Co.: 5-VIII-1956 (1♂ NDSU). McKenzie Co.: 146-98-16-110, 6-VII-1977, L. A. Schutz & J. W. Smith (1♀ NDSU), 21-VII-1976, Lago & Kurtz (1♂ NDSU), 25-VIII-1976 (1♂ NDSU). Mercer Co.: Stanton Area, 1-VII-1972, A. C. F. Hung (1♀ UNDG), 4-6-VIII-1972, A. C. F. Hung (1♀ UNDG); Sec. 22, T144N, R84W, 15-VIII-1972, P. B. Kannowski (1♂ UNDG). Morton Co.: 19-VIII-1958, R. L. Post (1♀ NDSU). Nelson Co.: 2-VII-1958 (1♀ NDSU). Oliver Co.: Sec. 6, T143N, R83W, 16-VIII-1972, P. B. Kannowski (1♀ DARC). Ransom Co.: Lisbon, 1-VII-1983, J. Julian (1♀ NDSU), 28-VII-1982, J. Julian, on leafy spurge (1♂ NDSU); McLeod, 19-V-1973, D. Scott (1♀ NDSU); 4mi. NW of McLeod, T134N, R53W, Sec. 6, 18-VI-1992, D. A. Rider (1♀ DARC); 7mi. SE Sheldon, 12-VII-1990, J. R. Powers (1♀ DARC). Richland Co.: Ekre Grassld Pres., 3-IX-2009, D. A. Rider (1♂ 2♀♀ DARC), Mirror Pool, 2-VII-1979, J. W. Hoganson (1♀ NDSU); Oak Savannah, T135N, R52WW Sec. 10, 11-VI-1996, L. DeCock (1♀ NDSU); Hwy 27, 5mi. W of Hwy 18, 5-IX-1979, E. U. Balsbaugh (1♀ NDSU); SNG, Horse trail head, 46.52N, 96.21W, 20-VII-2012, D. A. Rider (1♀ DARC); T136N, 61W Sec. 32, 23-VII-1993, G. Fauske (1♀ NDSU); 26-VII-1965, R. L. Post (1♀ NDSU). Slope Co.: Burning Coal Vein St. Pk., 23-VI-1965, L. Grochowski (1♀ NDSU), 1-VII-1965, L. Grochowski (1♀ NDSU), 10-VIII-1993, D. A. Rider (1♀ DARC); Chalky Buttes, 5-VI-1971, T. L. McCabe (1♂ NDSU); 136-102-14-200, 3-VII-1977, L. A. Schutz & J. W. Smith (1♂ NDSU), 29-VII- 1976, Lago & Kurtz (1♂ 4♀♀ NDSU), 8-VIII-1976 (2♂♂ 2♀♀ NDSU), 8-IX-1976, Lago & Kurtz (1♀ NDSU); 11-VII-1986, K. Larsen (1♀ MISU). Stutsman Co.: Jamestown, 7-VIII-1962, J. G. & B. L. Rozen (1♂ DARC). Ward Co.: Minot, 22-IV, Kluver (1♂ MISU), 13-V-1936, C. Bonness (1♂ MISU); SE Minot, 22-V- 1994, G. Hanley (1♀ MISU), 23-V-1994, G. Hanley (1♀ MISU); 6-V-1967, G. Eltz (1♀ MISU). Williams Co.: Williston, 23-VIII-1987, M. J. Weiss (1♀ NDSU).

Cosmopepla Stål, 1867 Members of this New World genus are relatively easy to identify by their small size, orbicular shape, and characteristic coloration. Most species are black with distinct dorsal markings in yellows, oranges, and/or reds (there are a few species in the western U.S. that are somewhat more drab, and as such appear quite similar to the Old World genus Eysarcoris). The genus has been revised fairly recently (McDonald 1986). There are presently eight species in the genus, all except one occurs in the western or southwestern U.S. extending into Mexico and Central America. Only one species occurs in North Dakota. Cosmopepla lintneriana Kirkaldy, 1909 - The name of this species has had a confusing taxonomic history which was discussed in length by Rider and Rolston (1995). In short, the original name carnifex Fabricius, 1798, is preoccupied; the next available synonym was thought to be bimaculata Thomas, 1865, but it is also preoccupied. Kirkaldy (1909) proposed lintneriana as a replace- ment name; this was not accepted by many workers. As such, much of the early literature pertaining to this species is under the name C. carnifex and most of the more recent literature is under C. bimaculata. Cosmopepla lintneriana is the correct name, however. This species is relatively easy to identify by the small orbicular shape, the black coloration, and the reddish dorsal markings including the pair of red spots near the scutellar apex, and a reddish cross on the pronotum. McPherson (1982, as C. bimaculata) gave the known range as Nova Scotia and New England west to British Columbia and Washington, and 336 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 south to Georgia, Texas, and Mexico. Uhler (1878, as C. carnifex) recorded this species from two North Dakota counties: Pembina Co.: near Pembina, June, and Rollette Co.: Turtle Mountains, July 24; Hussey (1922, as C. bimaculata) also recorded it from Rollette Co.: Turtle Mountains, T. H. Hubbell, taken on willow in a moist meadow. Lugger (1900) recorded this species (as C. carnifex) from Minnesota. The behavior, biology, and life history of this species has been studied (McDonald 1968b, Fish and Alcock 1973, McPherson 1976b, McPherson and Tecic 1997). Specimens examined: Barnes Co.: Bald Hill Dam, 7-VI-1978, E. U. Balsbaugh (2♂♂ NDSU); Valley City, 10-VI-1993, K. Mundal (2♂♂ DARC, 2♂♂ NDSU). Cass Co.: Fargo, 10-VI-1923, W. Baker (1♀ NDSU), 12-VII-1922, R. L. Webster (1♀ NDSU); N Fargo, Riverwood Pk, 46.945N, 96.802W, 19-VII- 2012, D. A. Rider (1♂ DARC); NE Fargo, 19-VI-2012, D. A. Rider (1♀ DARC); s. Fargo, 76 Ave S, Chrisan Edition, 46.766N, 96.780W, 5-VI-2012, D. A. Rider (1♀ DARC); Fargo, NDSU campus, 25-VI-2012, D. A. Rider (1♀ DARC); Tower City, 27-VI-1906, MWR (1♂ DARC, 1♂ 1♀ NDSU), 12-VII-1906 (1♀ DARC); Wild Rice R. at Red Riv., 46.758N, 96.787W, 1-VIII-2012, D. Rider (5♂♂ 2♀♀ DARC), 13-VIII-2012 (2♂♂ 1♀ DARC); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (3♂♂ 5♀♀ DARC); 17-III-1963, R. Poole (1♀ NDSU), 16-VI-1966 (1♀ NDSU), 24-VII-1968, J. Johnson (1♂ 3♀♀ NDSU), 9-VIII-1968, J. Johnson (1♂ NDSU). Cavalier Co.: Clyde, 20-IX-1923, W. Baker (1♂ NDSU); Langdon Farm, 7-IX-2002, D. Nowatzki (1♂ NDSU). Divide Co.: vicinity of Crosby, 13-IX-1987, M. Buck (1♂ MISU). Grand Forks Co.: Air Force Base Property, McKinock Twp., T152N, R53W, Sec. 27, 16-IX-1970 (2♀♀ UNDG); Forest R. Valley, UNDG Biol. Stn., 1mi. N Inkster, 9-VII-1976 (1♂ UNDG), 16-VII-1977 (1♂ UNDG), 30-IX-1970, D. Shouse (1♀ UNDG); Grand Forks, 24- VI-1953 (1♂ UNDG), 25-VI-1917, P. W. Fattig (1♂ 1♀ 1? NDSU), 26-VII-1953 (1♀ UNDG), 26-VII-1966 (1♀ UNDG), 29-IX-1945 (1♂ 1♀ UNDG), 2-X-1939 (1♀ UNDG), 4-X-1945 (1♀ UNDG), 13-X-1947 (1♀ UNDG); Grand Forks, 706 Reeves Dr., 20-IX-1982 (1♀ UNDG); Kelly, 1-X-1927, E. & G. Wheeler (1♂ UNDG); Northwood, 15-VI-1964, R. L. Post (8♂♂ 13♀♀ NDSU); Oakville Prairie, T151N, R52W, Sec. 16, 10-IX-1980 (1♂ 3♀♀ UNDG), 24-IX-1976 (1♀ UNDG), 10-X-1976 (1♂ UNDG); State Park near Arvilla, 30-VII-1966 (1♀ UNDG), 4-X- 1947 (2♂♂ UNDG); Turtle R., 4.8mi. W, 2mi. N Emerado, T152N, R53W, Sec. 20&29, 25-VI-1979 (1♀ UNDG), 9-VIII-1977 (1♂ UNDG), 8-IX-1975, A. Glass (1♀ UNDG), 8-IX-1980 (1♀ UNDG), 11-IX-1976 (1♀ UNDG), 12-IX-1977 (1♂ 1♀ UNDG), 16-IX-1984 (1♀ UNDG), 17-IX-1984 (1♀ UNDG), 18-IX-1976 (2♂♂ 3♀♀ UNDG), 20-IX-1978 (1♂ 1♀ UNDG); Turtle River State Park, 28-VI-1965, S. Kraft (1♀ UNDG), 4-IX-1976 (1♂ UNDG), 23-IX-1978 (2♂♂ UNDG); T151N, R50W, Sec. 6, 28-VII-1966 (1♀ UNDG); 9-IX-2008, T. Fettig (1♀ UNDG), 27-IX- 2008, C. Black (1♀ UNDG). Griggs Co.: Cooperstown, 30-VII-1964, R. J. Sauer (2♂♂ NDSU). McKenzie Co.: 1-VIII-2008, J. Beard (1♀ UNDG). Ransom Co.: Fort Ransom St. Pk., 13-VII-2002, D. A. Rider (4♂♂ 1♀ DARC); 2mi. SW McLeod, T134N, R53W, SW35, 9-VI-1992, D. A. Rider (1♂ 1♀ DARC), 8-VII- 1992, D. Cuthrell, T. Madson, at light (2♂♂ 1♀ NDSU); 4mi. NW of McCleod, T134N R53W Sec 6, 18-VI-1992, D. A. Rider (1♂ 6♀♀ DARC); Mirror Pool, 10mi. SE Sheldon, 12-VI-1981, J. Smith (1♀ NDSU); Mirror Pool WMA, 46°32.348N, 97°17.475W, 27-VI-2012, D. A. Rider (2♂♂ 6♀♀ DARC), 30-VIII-2006, W. Clark (1♀ NDSU); 7 mi. SE Sheldon, 23-V-1985, J. R. Powers (6♂♂ 3♀♀ DARC, 2♂♂ 2♀♀ NDSU); Sheyenne National Grassland, T134N, R53W, Sec. 12, 11-VI-1992, D. Cuthrell (1♀ NDSU); SNG, Orchid Site A, T134N, R54W, SESE12, 29-VI- 1992, D. Cuthrell (2♂♂ 4♀♀ NDSU); SNG, Orchid Site D, T134N, R53W, SE05, 23-VI-1992, D. Cuthrell (1♀ NDSU); SNG, Viking Prairie, 15-IX-2006, K. A. Fox (1♀ NDSU); 6-VI-1963, R. L. Post (1♀ NDSU), 11-VII-1961, J. C. Larsen (1♂ 1♀ NDSU). Richland Co.: Ekre Grassland Pres., 3-IX-2009, D. A. Rider (3♀♀ DARC); Mirror Pool, 3-VI-1994, D. A. Rider (1♀ DARC), 2-VII-1979, J. W. Hoganson (1♀ NDSU), 5-VII-1979, D. Fischer (1♂ NDSU); SNG, Oak Savannah, 2012 THE GREAT LAKES ENTOMOLOGIST 337 T135N, R52W, Sec. 10NE, 14-IX-1996, L. Pederson (1♂ NDSU); SNG, Horse trail head, 46.523N, 97.201W, 28-V-2010, D. A. Rider (7♂♂ 5♀♀ DARC; 7♂♂ 4♀♀ NDSU), 27-VI-2012, D. A. Rider (3♂♂ DARC), 25-VIII-2009, D. A. Rider (1♂ 1♀ DARC); Verbena Site, T135N, R51W, 31, 5-VIII-1992, D. Cuthrell (2♂♂ 2♀♀ NDSU). Traill Co.: 27-VI-1967, R. L. Post (2♂♂ 1♀ NDSU). Ward Co.: Minot, 29-VI-2008, D. Markle (1♂ NDSU), 22-VII-1948, Knapp & Dahl (1♂ 1♀ NDSU); Nedrose Twp, 5-VI-1957, R. Nelson (1♂ MISU). Williams Co.: T-154- 96-31, D. A. Mundal, Euphorbia esula (1♂ NDSU).

Euschistus Dallas, 1851 This is a large, diverse genus with many species occurring in North America and the Neotropics. There are no recent keys that cover all the North American species, the most recent being that of Torre-Bueno (1939). In general, however, North American species of Euschistus can be separated from other genera by the crenulate to dentate anterolateral pronotal margins. McPherson and Ahmad (2012) compared the male genitalia among those species of Euschis- tus that occur in the midwestern United States. Five species occur in North Dakota, and can be identified by the following key.

Key to the species of Euschistus in North Dakota 1 Pronotum with distinct transhumeral pale band or callous; spiracular peritremes dark brown to black...... Euschistus ictericus (Linnaeus) – Transhumeral pale band usually lacking, not distinct, if faintly present, then spiracular peritremes pale, concolorous with abdominal venter...... 2 2(1) Juga distinctly longer than tylus...... 3 – Juga and tylus subequal in length, juga rarely slightly longer than tylus in which case humeral angles are distinctly angulate or spinose...... 4 3(2) Connexivum immaculate...... Euschistus latimarginatus Zimmer – Connexivum pale with dark brown spots or bands...... Euschistus servus euschistoides (Vollenhoven) 4(2) Pale brown, somewhat more elongate; humeral angles angulate to spinose; abdominal venter lacking medial black spots...... Euschistus variolarius (Palisot de Beauvois) – Dark brown to gray, somewhat broader; humeral angles rounded; abdominal venter usually with one to four medial black spots...... Euschistus tristigmus luridus Dallas Euschistus ictericus (Linnaeus, 1763) - This species is relatively easy to recognize by the pale transhumeral callus and the dark spiracular peritremes. McPherson (1982) summarized the known distribution of this species as occurring in the northern states and Canada as far west as Utah, and as far south as Florida, Oklahoma, and Texas. Neither Uhler (1878) or Hussey (1922) reported this species from North Dakota. Froeschner (1988c) did not list this species from North Dakota, or from any of the surrounding states or provinces (the closest records are from Colorado, Iowa, Illinois, and Wisconsin). McPherson and Paskewitz (1984a) described the immature stages and observed the adults ovipositing on sedges. Specimens Examined: MINNESOTA [New State Record]: Clay Co.: MSUM Buffalo River Science Center, 46°51′45″N, 96°28′41″W, 24-VIII-2006, W. Clark (1♂ DARC), 26-VIII-2004, S. Ganehiarachchi (1♀ NDSU), C. P. Johnson (1♀ DARC). Dakota Co.: Hastings, 20-VIII-1928, H. A. Scullen (1♂ DARC). Stearns Co.: St. Cloud, 1-VIII-1972 (1♀ DARC); St. Stephen, 18-IX-1968, R. J. Trobec (1♂ UNDG). NORTH DAKOTA [New State Record]: Cass Co.: Fargo, 338 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

10-VI-1923, W. Baker (1♀ NDSU); Fargo, NDSU campus, Ladd Hall, 1-X-2002, R. Bohl (1♀ NDSU); Wild Rice R. at Red Riv., 46.758N, 96.787W, 1-VIII-2012, D. Rider (1♀ DARC); IX-1958 (1♂ NDSU). Grand Forks Co.: 28-VIII-2008, T. Campbell (1♂ UNDG); State Park near Arvilla, 4-X-1947 (1♂ UNDG). Ransom Co.: SNG, A-Annex, 46.441N, 97.384W, 6-IX-2001, D. Rider (1♂ DARC). Euschistus latimarginatus Zimmer, 1910 - This species can be recognized by the elongate juga and the immaculate connexiva. Euschistus latimarginatus was originally described from Nebraska, and was later recorded from South Dakota (Harris, 1937). Van Duzee (1917) listed it also from Colorado (repeated by Torre-Bueno, 1939 and Froeschner 1988c), but the source of this record has not been located. From specimens at hand, this species appears to be much more widespread than previously thought, but it may be very local, occurring only on sand dune sites. Zimmer (1912) indicated that he collected this species on seed pods of yucca and opuntia; I have collected a single specimen (as a late instar which I reared to adult on green beans) on the seed pod of penstemon. Because of the apparent lack of published records, I have listed several new state records outside the scope of this work. Specimens Examined: MINNESOTA [New State Record]: Polk Co.: 2mi. S Fertile, 47°30.14N, 96°18.00W, 12-VII-2000, P. B. Beauzay (1♀ NDSU). NEBRASKA: Cherry Co.: Valentine, 10-VI-1982, Wolfe (1♂ DARC). Thomas Co.: Halsey, 12-VI-1971, K. Pruess, on yucca (5♂♂ 3♀♀ DARC). NORTH DA- KOTA [New State Record]: Ransom Co.: Sheyenne Natl. Grasslands, T135N, R53W, Sec. 27, 28-VII-2000, J. Maxwell, in sand hills nr Hudsonia spp. (1♂ NDSU). Richland Co.: Mirror Pool, 6-IX-1980, J. Thomas (1♂ NDSU); 1mi. E Mirror Pool, 24-VII-1993, coll. as nymph on penstamon (1♂ DARC). SOUTH DAKOTA: Bennett Co.: Martin, 3-IX-1924, H. C. Severin (1♂ DARC). TEXAS [New State Record]: El Paso Co.: Fabens, IX-1940, S. E. Jones (1♂ DARC). Ward Co.:Monahans Sand Hills, 28-VII-1973, W. E. & C. A. Triplehorn (1♂ DARC); 3mi. E. Monahans, 19-VIII-1983, E. G. & M. A. Riley (1♂ DARC). UTAH [New State Record]: Zion Natl Park (1♂ DARC). Euschistus servus euschistoides (Vollenhoven, 1868) - In North Dakota, this species is easy to recognize by the elongate juga and the maculate connexiva. The nominate subspecies is more difficult to separate from other Euschistus species as the juga and tylus are subequal in length; this subspecies occurs in the southern half of the U.S. There is some question whether euschistoides is a valid subspecies as the boundary between the two subspecies is rather broad, and the main character separating the two subspecies is more or less clinal through this boundary area. McPherson (1982) stated that the boundary between the two subspecies was not clear except for his work in Illi- nois and some work in Virginia; he gave the range for this subspecies as simply across the northern part of the continent. Froeschner (1988c) did not list this species from North Dakota or Minnesota even though Hussey (1922) provided two North Dakota records: Ramsey Co.: Devils Lake, T. H. Hubbell, and Slope Co.: Amidon, T. H. Hubbell. I originally had thought that this species had not actually been recorded from Minnesota, but upon closer inspection, I discovered that the record of E. variolarius (Lugger 1900) is actually a misidentification of this species. In northern regions, E. variolarius has the humeral angles spinose; the specimen in Lugger’s illustration distinctly has rounded humeral angles. This species appears to be quite common, and probably occurs statewide in both North Dakota and Minnesota. Youther and McPherson (1975) studied the mating behavior of this species in the laboratory. This common species has been the subject of numerous citations concerning its biology and life history (see McPherson 1982 for citations). Specimens Examined: NORTH DAKOTA: Barnes Co.: Valley City, 10- VI-1993, K. Mundal (1♀ DARC); T139, R58, Sec. 14, 18-IX-2004, P. S. Burange (1♂ 1♀ NDSU). Benson Co.: 8-IX-1958, S. Wilson (1♂ NDSU). Billings Co.: 2012 THE GREAT LAKES ENTOMOLOGIST 339 Roosevelt Natl. Park, Medora, 24-VI-2002, F. Antwi (1♂ NDSU). Burleigh Co.: 12-VI-2008, B. Axtman (1♂ UNDG), 17-VII-2008, H. Dassinger (1♀ UNDG). Cass Co.: Absaraka, 19-IX-2008, L. Grindberg (1♂ 1♀ NDSU); Fargo, 24-VIII- 1923, W. Baker (1♂ NDSU), 29-VIII-1917, P. W. Fattig (1♀ NDSU), 7-IX-2004, L. Kuster (1♂ NDSU); Fargo, NDSU campus, 9-IV-2008, R. DeGreeff (1♂ NDSU), 11-IX-2008 (1♂ NDSU); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (1♂ DARC), 1-VIII-2012, D. A. Rider (1♀ DARC); 5-VIII- 1960 (1♂ NDSU), 9-IX-2003, R. Wagner (1♂ NDSU). Cavalier Co.: Langdon Farm, 7-IX-2002, D. Nowatski (1♂ NDSU); 5mi. E Langdon, 25-VIII-2007, J. Balsdon (1♂ NDSU). Dunn Co.: 5mi. W Killdeer, 26-V-1977, E. U. Balsbaugh (1♀ NDSU); 21-VI-1977, L. A. Schutz & J. W. Smith (1♂ NDSU), 13-VII-1976, Lago & Kurtz (1♀ NDSU), 20-VII-1977, L. A. Schutz & J. W. Smith (1♀ NDSU), 2-VIII-1977, L. A. Schutz & J. W. Smith (1♂ 1♀ NDSU), 25-VIII-1976 (2♂♂ 2♀♀ NDSU). Golden Valley Co.: D. A. Mundal, Eurphorbia esula (1♂ 2♀♀ NDSU). Grand Forks Co.: 5mi. N of Arvilla, 7-IX-2002, K. Benson (1♂ NDSU); State Park near Arvilla, 4-X-1947 (1♀ UNDG); Forest R. Biol. Stn., 1mi. N Inkster, 9-VII-1976 (2♀♀ UNDG); Grand Forks, 11-IX-1992 (1♀ UNDG), 4-X-1983 (1♂ UNDG); Grand Forks Industrial Park, 1-IX-2011, G. Machart, Lindgren trap baited with Ips lure (1♂ NDSU); Grand Forks KOA, 1-IX-2011, G. Machart, Lindgren trap baited with Manuka oil (1♀ NDSU); Northwood, 1-VI-1956, R. C. & R. L. Post (1♀ NDSU); Oakville Prairie, T151N, R52W, Sec. 16, 22-VI-2012, D. A. Rider (1♂ 1♀ DARC), 14-VIII-2009, D. A. Rider (1♂ DARC), 28-VIII-2008, B. Stargardt (1♀ UNDG), 10-IX-1980 (1♂ UNDG), 15-IX-1975 (1♀ UNDG), 15-IX- 1979 (1♀ UNDG), 26-IX-1983 (1♂ UNDG); Turtle River State Park, 4-IX-1976 (1♂ 1♀ UNDG), 2-X-1992 (1♀ UNDG); Turtle R., 4.8mi. W, 2mi. N Emerado, T152N, R53W, Secs. 20 & 29, 12-VI-1977 (1♀ UNDG), 9-VIII-1979 (1♂ UNDG). 1-IX-1982 (1♀ UNDG), 2-IX-1992 (1♂ 1♀ UNDG), 6-IX-1980 (5♂♂ 2♀♀ UNDG), 6-IX-1992 (1♂ UNDG), 8-IX-1975, A. Glass (1♂ 1♀ UNDG), 8-IX-1982, R. Kreil (2♂♂ 2♀♀ UNDG), 9-IX-1975 (1♀ UNDG), 9-IX-1982 (1♂ UNDG), 10-IX-1980 (2♀♀ UNDG), 17-IX-1984 (1♀ UNDG), 18-IX-1974 (1♀ UNDG), 19-IX-1984 (1♀ UNDG); UND campus, 19-X-1975 (1♂ UNDG); T151N, R50W, Sec. 26, 9-X-1969 (1♂ UNDG); 7-VIII-2008, J. Hughes (1♂ UNDG), 9-IX-2008, H. Dassinger (1♂ UNDG), 9-IX-2008, T. Fettig (1♂ UNDG), 12-IX-2008, P. Dorgan (1♂ UNDG), 12-IX-2008, C. Black (1♀ UNDG), 18-IX-2008, M. Michelson (1♀ UNDG), 30-IX- 2008, N. Hoagland (2♀♀ UNDG), 4-XI-2008, N. Hoagland (1♂ UNDG), 13-XI- 2008, B. Stargardt (1♀ UNDG). Marshall Co.: 4mi. W Argyle, 27-IX-2008, K. Rivard (1♀ NDSU). McHenry Co.: T156N, R78W, Sec. 36, 22-VI-1967, J. Knud- son (1♀ NDSU). McKenzie Co.: Th. Roos. M. Prk., 3-VIII-1965, W. Kotchman (1♂ NDSU). Mercer Co.: Stanton Area, 1-VII-1972, A. C. F. Hung (1♀ UNDG); T144N, R84W, Sec. 22, 6-VII-1972, A. C. F. Hung (1♂ 4♀♀ UNDG), T144N, R84W, Sec. 28, 18-20-VII-1972, A. C. F. Hung (1♂ UNDG); T144N, R85W, Sec. 9, 18-20-VII-1972, A. C. F. Hung (1♂ UNDG), 15-VIII-1972, P. B. Kannowski (1♂ UNDG). Mountrail Co.: Van Hook, 5-VII-1982 (1♀ NDSU). Oliver Co.: 2mi. E. of Hensler, 20-VII-1978, L. A. Schutz & J. W. Smith (1♀ NDSU); T143N, R83W, Sec. 6, 16-VIII-1972, P. B. Kannowski (1♀ UNDG). Pembina Co.: Crystal, 22-IX-2004, P. S. Burange (1♂ NDSU); 2mi. NE Crystal, 13-IX-2003, A. Moquist (1♀ NDSU); Icelandic State Park, 27-VIII-1981, E. U. Balsbaugh (1♀ NDSU). Ramsey Co.: Lawton, 22-IX-2004, R. Miller (1♀ NDSU). Ransom Co.: Fort Ransom St. Pk, 13-VII-2002, D. A. Rider (1♀ DARC); Lisbon, 1983, J. Julian, on leafy spurge (2♂♂ 1♀ NDSU); 2mi. SE McLeod, T134N, R53W, Sec. 35, 8-VII-1992, D. Cuthrell, T. Madson, at light (1♀ NDSU); Mirror Pool WMA, 27-VI-2012, D. A. Rider (1♀ DARC), 30-VIII-2007, D. A. Rider (3♂♂ 1♀ DARC), 4-IX-2009, D. A. Rider (1♂ DARC); 7mi. SE Sheldon, 19-V-1992, J. R. Powers (1♀ DARC), 23-V-1985, J. R. Powers (3♂♂ 1♀ DARC), 5-IX-1979, J. W. Smith (1♂ NDSU); SNG, A-Annex, 46.441N, 97.384W, 6-IX-2001, D. Rider (1♀ DARC). Richland Co.: 4mi. W Barney, 14-VI-1984, M. McLeod (1♂ NDSU); Colfax, 17- IX-2000, A. Hackey (1♂ NDSU); Ekre Grassland Preserve, T136N, R51W, Sec. 32, 28-V-2010, D. A. Rider (2♀♀ DARC), 28-VII-1993, D. L. Cuthrell (1♀ DARC), 340 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

19-VIII-2011, D. Rider, G. Fauske, Lindgren trap (♀ NDSU), 28-VIII-2008, D. A. Rider (2♂♂ DARC), 31-VIII-2011, D. Rider, G. Fauske, Lindgren trap (1♂ NDSU), 3-IX-2009, D. A. Rider (1♂ 4♀♀ DARC), 6-IX-1994, M. Ellefson (2♀♀ NDSU), 6-IX-1994, D. A. Rider (1♀ DARC), 25-IX-1994, T. Nowatzki (1♂ 1♀ NDSU); SW of Kindred, 11-IX-1993, J. Franger (1♀ NDSU); McLeod, T133N, 52W, Sec. 6, 18-IX-1979, E. U. Balsbaugh (1♂ NDSU); 1mi. SE McLeod, 3-VI- 1985, J. R. Powers (1♀ DARC); Mirror Pool, T135N, R52W, Sec. 8, 3-VIII-2004, S. Ganehiarachchi (1♂ 2♀♀ NDSU), 26-VIII-1994, T. Nowatzki (3♂♂ NDSU), 29-VIII-1996, D. A. Rider (2♂♂ 1♀ DARC), 3-IX-1992, J. A. Nelson (1♀ NDSU), 4-IX-1998, D. A. Rider (1♂ DARC), 11-IX-1993, B. Clapper (1♂ NDSU), 18-IX- 1994, D. A. Rider (1♀ DARC), 20-IX-1994, W. G. Schroeder (1♂ NDSU); 1mi. E Mirror Pool, 25-VI-1993, D. L. Cuthrell (1♀ DARC), 28-VII-1993, D. Cuthrell (1♂ NDSU); 2mi. NE Mirror Pool, 28-VI-1993, G. Fauske (1♀ NDSU); SNG, Oak Savannah Area, 11-VI-1996, L. DeCock (1♀ NDSU), 18-VI-2000, D. A. Rider (1♂ DARC), 26-VI-1996, L. DeCock (1♀ NDSU), 7-VIII-1993, D. Cuthrell (1♂ 1♀ NDSU), 17-IX-1996, P. Anderson (2♂♂ NDSU); SNG, Horse trail head, 46°31.409N, 97°12.194W, 25-VIII-2009, D. A. Rider (3♂♂ 1♀ DARC), 4-IX-2009, D. A. Rider (2♂♂ 3♀♀ DARC), 9-IX-2008, D. A. Rider (1♂ DARC); SNG Horse trail head, 46.52N, 96.21W, 20-VII-2012, D. A. Rider, col. on Penstamon as last instar nymph (1♂ DARC); Walcott, 10-IX-2004, C. Juricek (1♂ NDSU); Walcott Dunes, 26-VI-1975, P. K. Lago (1♀ NDSU); 11mi. W. Walcott, 23-V-1988, J. R. Powers (1♂ 2♀♀ DARC); 4-H Camp, 2-VII-1964 (1♂ NDSU); T135N, R52W, S4- SW1/4, 26-IX-1984, R. Kruger (1♂ NDSU); 6-VIII-1958, S. Wilson (1♀ NDSU). Rollette Co.: 4-VI-1958, Kotchman (1♂ NDSU). Sargent Co.: 3.5mi. N Crete, 5-IX-1982 (1♀ UNDG). Slope Co.: Chalky Buttes, 5-VI-1973, T. L. McCabe (1♀ NDSU). Stearns Co.: St. Stephen, 28-IX-1968 (1♂ UNDG). Stutsman Co.: Chase Lake, 13-VI-1975, P. K. & B. A. Lago (1♀ NDSU). Walsh Co.: 10mi. W of Park River, 11-VI-1981, E. U. Balsbaugh, on Carduus nutans (3♂♂ 3♀♀ NDSU); South Branch of Park River, NE of Adams, 20-VIII-1970, J. Singleton (1♂ UNDG); 20-VII-2008, E. Oldenburg (1♂ UNDG), 23-VII-2008, M. Flom (1♀ UNDG), 16-VIII-1958 (1♀ NDSU). Ward Co.: Minot 15-VI-1936, R. Neubauer (1♂ MISU), 10-VII-2002, G. Hanley (2♂♂ 1♀ MISU); NW Minot, 6-VI-1994, G. Hanley (1♀ MISU), SE Minot, 22-VI-2005, G. Hanley (1♀ MISU); Nedrose Twp, 12-VI-1957, R. Nelson (1♀ MISU). Euschistus tristigmus luridus Dallas, 1851 - This species is usually easy to identify, because the abdominal venter has from one to four black spots along its midline. In a few specimens, these spots are obsolete, but they can still be identified by their smaller size, the rounded humeral angles, and the dorsum being somewhat darker brown to gray, contrasting more sharply with the pale venter. Although this species can be caught in old field situations, it tends to occur more in forested areas than the other North Dakotan Euschistus species. Similar to E. servus euschistoides, this species has been divided into two subspecies. The northern subspecies, E. t. luridus, has the humeral angles rounded, whereas the nominate subspecies (southeastern U.S.) has the humeral angles acute to spinose. Apparently the boundary between these two forms is much more narrow than in E. t. euschistoides; McPherson (1976a) gave the dividing line as around latitude 41° N and gave (1982) the distribution north of that latitude from Nova Scotia, Québec, and New England west to Washington. Only the luridus subspecies occurs in North Dakota. Hussey (1922) provided two North Dakota records of this species: Eddy Co.: Sheyenne River, about 3 miles S of Warwick, 8 August 1920, T. H. Hubbell, and Ramsey Co.: Devils Lake, 28 July, T. H. Hubbell, undergrowth in an oak woods. It appears that this species is common throughout the eastern half of the state, but seems to be absent in the far western counties. The life history of this species has been studied (McPherson 1975), and this species has been reared in the laboratory (McPherson 1971b). 2012 THE GREAT LAKES ENTOMOLOGIST 341

Specimens Examined: NORTH DAKOTA: Barnes Co.: Valley City 10-VI-1993, K. Mundal (1♂ DARC); 13-IX-1946, R. L. Post (1♂ NDSU). Cass Co.: Fargo, VII-1957, R. C. & G. J. Post (1♀ NDSU), 31-VIII-2004, N. Hillen (1♀ NDSU); Fargo, NDSU campus, 3-IX-2002, R. Holthusen (1♂ NDSU); S Fargo, 76 Ave. S., Chrisan Edition, 46.766N, 96.780W, 8-VI-2012, D. A. Rider (1♂ 1♀ DARC); 17-VI-1958, S. Wilson (1♂ NDSU). Cavalier Co.: Langdon Farm, 7-IX-2002, D. Nowatzki (1♂ NDSU). Grand Forks Co.: Air Force Base Property, McKinock Twp, T152N R53W, Sec. 27, 21-IX-1974 (1♀ UNDG); Grand Forks, 20-VIII-1928 (1♀ UNDG), 26-IX-1952 (1♂ UNDG), 3-X-1941 (1♂ UNDG); Northwood, 15-VI-1964, R. L. Post (1♀ NDSU); Oakville Prairie, Oakville Twp, Sec. 9, 2-X-1958 (1♂ UNDG); Oakville Prairie, Sec. 16, 47°53.403N, 97°18.855W, 22-VI-2012, D. A. Rider (1♂ 5♀♀ DARC), 21-VIII-2009, D. A. Rider (1♂ DARC), 10-IX-1980 (1♀ UNDG); Turtle River, 4.8mi. W, 2mi. N. Emerado, T152N, R53W, Sec. 20 & 29, 8-IX-1975, A. Glass (1♀ UNDG), 10-IX-1980 (1♂ UNDG), 17-IX- 1984 (2♂♂ 1? UNDG), 18-IX-1974 (1♂ UNDG), 19-IX-1984 (1♂ 1♀ UNDG), 25-IX-1965 (1♀ UNDG); Turtle River State Park, 8-VII-1940 (1♂ UNDG), 14-IX-1975 (2♂♂ UNDG), 18-IX-1965 (1♀ UNDG), 19-IX-1966 (1♂ UNDG), 25- IX-1937 (1♀ UNDG), 27-IX-1969, R. Becker (1♀ UNDG), 4-X-1947 (1♂ UNDG); UNDG Biology Area, Forest R. Valley, Inkster, 18-VI-1958 (1♀ UNDG), 18-VI- 1970, R. Lamp (1♀ UNDG), 25-VI-1970, G. Ruonala (1♂ UNDG), 9-VII-1976 (1♀ UNDG), 31-VIII-1966 (2♀♀ UNDG), 29-IX-1980 (1♂ UNDG), 5-X-1969 (1♂ UNDG), 9-X-1965 (1♀ UNDG); 25-VIII-2008, B. Roth (1♀ UNDG), 10-IX-2008, J. Morseth (1♀ UNDG). Mercer Co.: T144N, R84W, Sec. 22, 6-VII-1972, A. C. F. Hung (2♂♂ 3♀♀ UNDG), 4-6-VIII-1972, A. C. F. Hung (1♀ UNDG), 15-VIII- 1972, P. B. Kannowski (2♀♀ UNDG); Stanton area, 1-VII-1972, A. C. F. Hung (1♀ UNDG). Morton Co.: Mandan, VII-1965, J. W. Goertel (1♂ NDSU); 12- VIII-2008, N. Bundermann (1♂ UNDG). Pembina Co.: 4mi. W of St. Thomas, 14-VI-1979, J. Pontius, window pane trap (1♂ NDSU). Ramsey Co.: Lawton, 17-IX-2004, M. Duppong (1♀ NDSU). Ransom Co.: Lisbon, 1983, J. Julian (1♀ NDSU); Mirror Pool WMA, 46°32.348N, 97°17.475W, 27-VI-2012, D. A. Rider (3♀♀ DARC), 18-VII-2002, L. Tobias (1♀ DARC), 24-VII-1979, A. C. Ashworth (1♂ NDSU), 30-VIII-2007, D. A. Rider (3♂♂ DARC); SNG, A-Annex, 46.441N, 97.384W, 6-IX-2001, D. Rider (1♂ DARC). Richland Co.: Ekre Grassland Pre- serve, T136N, R51W, Sec. 32, D. Pitzen (1♂ 2♀♀ NDSU), 1-V-2011, D. Rider, G. Fauske, Lindgren trap (8♂♂ 4♀♀ NDSU), 28-V-2010, D. A. Rider (2♂♂ 2♀♀ DARC), 8-VII-2011, D. Rider, G. Fauske, Lindgren trap (1♂ 5♀♀ NDSU), 21- VII-2011, D. Rider, G. Fauske, Lindgren trap (1♂ 5♀♀ NDSU), 28-VII-1993, D. L. Cuthrell (2♀♀ DARC), 3-IX-2009, D. A. Rider (3♂♂ 8♀♀ DARC), 11-IX-1993, M. J. Dietrich (1♀ NDSU); Mirror Pool, T135N, R52W, Sec. 8, 27-VI-1992, D. Rider (1♀ DARC), 2-VII-1979, J. W. Hoganson (1♀ NDSU), 18-VII-1992, D. A. Rider (1♀ NDSU), 10-VIII-1994, J. L. Jyoti (1♂ 2♀♀ NDSU), 29-VIII-1996, D. Rider (4♂♂ 2♀♀ DARC), 4-IX-1998, D. A. Rider (3♂♂ 1♀ DARC), 5-IX-1994, T. Nowatzki (1♂ NDSU), 11-IX-1974, P. K. Lago (1♀ NDSU); 1mi. E Mirror Pool, 25-VI-1993, D. L. Cuthrell (1♂ DARC); 1.5mi. NE Mirror Pool, 16-VI-1994, D. A. Rider (1♀ DARC), 27-VII-1993, D. L. Cuthrell (1♀ DARC); SNG, Horse Trail head, 46°31.409N, 97°12.194W, 27-VI-2012, D. A. Rider (1♂ 1♀ DARC), 4-IX- 2009, D. A. Rider (1♀ DARC); SNG, Oak Savannah, 18-VI-2000, D. A. Rider (1♀ DARC). Rollette Co.: Sec. 20, Twp. 163, R. 72, 9-VI-1967, J. Knudson (1♀ UNDG); 4-VI-1968, Kotchman (1♀ NDSU). Sargent Co.: Shuman Twp, NW Sec. 3, 12-IX-1976 (1♂ UNDG). Stutsman Co.: 27-IX-1958, R. L. Post (1♀ NDSU). Traill Co.: 27-VI-1967, R. L. Post (1♀ NDSU). Walsh Co.: S. Branch, Park River, NE of Adams, 19-VIII-1970, D. Shouse (1♂ UNDG), 21-VIII-1970, J. Singleton (1♂ UNDG). Euschistus variolarius (Palisot de Beauvois, 1817) - This species is commonly referred to as the onespotted stink bug, the one spot being a black medial spot at the base of the pygophore in males. Euschistus variolarius is a common, widespread species, often confused with E. servus. Male specimens 342 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 are easy to separate because the black spot on the pygophore (referred to above) is lacking in E. servus. In North Dakota, the two are readily separated by the juga (elongate in E. servus, not so in E. variolarius) and the humeral angles (rounded in E. servus, angulate to spinose in E. variolarius). One should note that these characters work only in northern latitudes; E. servus is quite variable in more southern areas. McPherson (1982) gave the distribution of this species as Québec, Ontario, and New England south to Florida, and west to British Columbia, Idaho, and Utah. Neither Uhler (1878) nor Hussey (1922) recorded collecting specimens of this species from North Dakota; also Froeschner (1988c) did not list this species from North Dakota or surrounding states. Lugger (1900) recorded this species from Minnesota, but this record is a misidentification of E. servus. The North Dakota distribution of this species is very interesting. There are only two North Dakota records of E. variolarius prior to 1993, one in Richland County in 1958 and the other from Grand Forks County in 1976. Since 2000, however, this species has been collected fairly frequently, especially in the southeastern part of the state, but there is a record as far north and west as Minot in Ward County (2002). The biology of this species has been studied (Parish 1934), and Munyaneza and McPherson (1994) described the life cycle of this species in the midwest. Specimens Examined: NORTH DAKOTA [New State Record]: Cass Co.: Fargo, 46°54′55″N, 96°47′55″W, 21-VII-2002, L. M. Tobias, UV light trap (1♂ DARC); Fargo, 19th Ave. SW, 24-IX-2002, E. Delahoyde (1♀ NDSU); s. Fargo, 76 Ave S., Chrisan Edition, 46.766N, 96.780W, 5-VI-2012, D. A. Rider (1♂ DARC); Fargo, NDSU campus, 21-V-2012, D. A. Rider (2♂♂ DARC), 25-VI-2012, D. A. Rider (1♂ 2♀♀ DARC), 6-IX-2000, G. Goenner (1♀ NDSU), 6-IX-2001, D. A. Rider (1♂ 1♀ DARC), 7-IX-2002, F. Antwi (1♀ NDSU), 9-XI-2008, J. Ell (1♀ NDSU); s. Fargo, 10-IX-2002, S. Koudela (1♀ NDSU); Fargo, Edgewood Golf Course, 2-X-2002, J. Prust (1♂ NDSU); Wild Rice R. at Red Riv., 46.758N, 96.787W, 1-VIII-2012, D. Rider (4♂♂ 1♀ DARC), 13-VIII-2001, D. A. Rider (1♂ DARC); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 13-VIII-2012, D. A. Rider (1♂ DARC). Grand Forks Co.: Oakville Prairie, Sec. 16, 47°53.403N, 97°18.855W, 22-VI-2012, D. A. Rider (1♂ 2♀♀ DARC); Turtle River, 4.8mi. W, 2mi. N Emerado, T152N, R53W, Sec. 20 & 29, 15-IX-1976 (1♂ UNDG); 28-VIII- 2008, D. Johns (1♂ UNDG). Griggs Co.: 1mi. E Hannaford, 8-XI-2007, K. D. Haugen (1♂ NDSU). LaMoure Co.: 3mi. W LaMoure, 28-IX-2002, C. Dathe (1♂ NDSU). Ransom Co.: Mirror Pool WMA, 27-VI-2012, D. A. Rider (2♂♂ 5♀♀ DARC), 30-VIII-2006, C. P. Johnson (1♀ NDSU), 30-VIII-2007, D. A. Rider (1♀ DARC), 7-IX-2000, C. L. Jorgensen (1♂ NDSU); SNG, A-Annex, 46.441N, 97.384W, 6-IX-2001, D. Rider (2♂♂ 1♀ DARC). Richland Co.: Ekre Grassland Preserve, 15-VII-2002, A. Joshi (1♂ 1♀ NDSU), 28-VII-1993, D. Cuthrell (1♂ DARC); Mirror Pool, 3-VIII-2004, S. Ganehiarachchi (1♂ NDSU); SNG, Horse- trail Head nr. Oak Savannah, 25-X-2007, D. A. Rider (1♂ DARC); SNG, Horse trail head, 46°31.409N, 97°12.194W, 27-VI-2012, D. A. Rider (2♂♂ DARC), 8-IX-2008, D. A. Rider (1♀ DARC); SNG, Horsetrail head, 46.52N, 96.21W, 20-VII-2012, D. A. Rider, col. on Penstamon as last instar nymph (1♀ DARC); 6-VIII-1958, S. Wilson (1♀ NDSU); SNG, Oak Savannah, West I Allotment, 3-IX-2002, F. Antwi (1♂ 1♀ NDSU), A. Joshi (1♀ NDSU). Ward Co.: Minot, 10-VII-2002, G. Hanley (2♀♀ MISU).

Holcostethus Fieber, 1860 This genus has had a confused taxonomic history, and there are still some existing problems. The taxonomic history has been given in detail in Belousova (2007) and Ribes et al. (2008). In recent times, nearly all species were placed in Holcostethus. Ribes and Schmitz (1992) proposed that those species in which the tylus is enclosed by the juga should be placed in Peribalus (Ribes and Schmitz erroneously used Dryocoris for this genus - see Dolling 1995, and Rider and 2012 THE GREAT LAKES ENTOMOLOGIST 343 Rolston 1995 for an explanation), whereas those with the tylus not enclosed by the juga belonged in Holcostethus. Rider and Rolston (1995) argued that in North American species (not studied by Ribes and Schmitz), this character was variable even within a species, and as such only one genus, Holcostethus, should be recognized. Both Belousova (2007) and Ribes et al. (2008) have again argued that both genera should be recognized, and now have added several characters from both the male and female genitalia to support their position. These authors still did not study New World species. I have examined specimens of most of the North American species and have conducted a quick survey of the New World literature (McDonald 1974, 1982), and conclude once again that in New World species, the jugal characters are variable, and simply do not work. The other characters, although not completely surveyed, seem to place most, if not all of our North American species in Peribalus. It seems best, however, to keep our species in Holcostethus until a thorough study of North American taxa can be made. The North American species were revised by McDonald (1974); he later provided an updated key (McDonald 1982). There are presently three species of Holcostethus known to occur in North Dakota. One other species, H. fulvipes, has been recorded from Manitoba (Maw et al. 2000). Key to species of Holcostethus that may occur in North Dakota 1 Abdominal venter dark brown with at most the margins outlined in yellow...... macdonaldi Rider and Rolston – Abdominal venter pale yellow to reddish brown, with or without dark markings...... 2 2(1) Reddish brown; abdominal venter with distinct black markings in zig-zag pattern; juga not meeting in front of tylus [not yet known from North Dakota]...... fulvipes (Ruckes) – Brown; abdominal venter without distinct pattern of dark markings.3 3(2) Connexiva pale with a dark transverse band along both anterior and posterior connexival margins, resulting in an alternating pale and dark pattern...... abbreviatus Uhler – Connexiva with narrow mesial margins dark, lateral margins pale, not interrupted by dark bands, no alternating pale and dark pattern. limbolarius (Stål) Holcostethus abbreviatus Uhler, 1872 - In this species, the tylus is distinctly enclosed by the juga, and the connexiva is distinctly banded, alternating dark and pale. McPherson (1982) gave the range of this species as Iowa northwest to British Columbia and Washington, west to Oregon and California, and southwest to New Mexico. Hussey (1922) provided two specific localities for North Dakota; they were Bottineau Co.: Bottineau, from herbage in a depression in a pasture, T. H. Hubbell, and Rollette Co.: Turtle Mountains, shrub-filled clearing, T. H. Hubbell. This species has also been recorded from Manitoba (Maw et al. 2000). Interestingly, it appears that this species has not been recorded from Minnesota yet. Specimens Examined: MINNESOTA [New State Record]: Clay Co.: CCL Trust Lands, 47°03′Lat. 96°26′Long., 1-IX-1997, P. Tinerella, pitfall trap (1♀ NDSU); Felton Prairie Complex, 3mi. E, 2mi. S Felton, 47°03.87′N, 96°25.64′W, 30-VII-1999, David, Jordan (1♀ NDSU). NORTH DAKOTA: Billings Co.: Sully Crk. St. Park, 10-VIII-1993, D. A. Rider (1♀ DARC). Cass Co.: Fargo, 10-VI-1923, W. Baker (1♀ NDSU); Fargo, NDSU campus, 27-IX-2008, A. Clyle (1♀ NDSU). Dunn Co.: Killdeer Mts., 28-VI-1965, L. Grochowski (1♀ NDSU); 3-VII-1966 (1♀ NDSU). Grand Forks Co.: Forest River Biol. Area, 1.8mi. W, 1mi. N Inkster, T154N, R55W, Sec. 11 & 14, 18-IX-1976 (1♀ UNDG); Oakville Praire, Oakville Twp., Sec. 9, 23-VI-1961 (1♀ UNDG). Mercer Co.: Stanton 344 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 area, 1-VII-1972, A. C. F. Hung (♀ UNDG); T144N, R84W, Sec. 22, 4-6-VIII- 1972, A. C. F. Hung (1♀ UNDG). Morton Co.: 6mi. E Glen Ullin, 1-VIII-1975, P. K. & B. A. Lago (1♀ NDSU). Ransom Co.: SNG, 1.5mi. W McLeod, T134N, R53W, Sec. 22, 4-VI-1992, D. A. Rider (1♀ DARC). Richland Co.: 1.5mi. NE Mirror Pool, 18-VII-1993, D. L. Cuthrell (1♀ NDSU). Slope Co.: Burning Coal Vein St. Pk., 1-VI-1965, W. Kotchman (1♀ NDSU), 6-VI-1973, T. L. McCabe (1♀ NDSU), 13-VII-1965, W. Kotchman (1♀ NDSU), 10-VIII-1993, D. A. Rider (1♀ DARC); Chalky Buttes, 7-VI-1965, L. Grochowski (1♀ NDSU). Stutsman Co.: Montpelier, 31-V-1972, M. Knutson (1♀ NDSU). Ward Co.: Minot, 9-V-1936, O. Hill (1♂ MISU), 19-V-1936, C. Johnson (1♂ MISU); SE Minot, 23-VI-2005, G. Hanley (1♀ MISU). Holcostethus limbolarius (Stål, 1872) - This is the most common and widespread species in North America. This species is most likely to be confused with P. abbreviatus, but the two are easily separated by the pattern of dark markings on the connexival segments. In P. abbreviatus, the connexiva has a distinct alternating pale and dark pattern, whereas in P. limbolarius, the lateral half of the connexiva is entirely pale, not interrupted with dark. McPherson (1982) gave the range of this species as Québec, Ontario, and New England west across the continent, and south and southwest to Georgia, Texas, New Mexico, and California. This species appears to be fairly common in North Dakota, but to date only a single literature record is known. Hussey (1922) recorded it from Slope Co.: Amidon, T. H. Hubbell. Interestingly, it appears that this species has not yet been documented for Minnesota; Froeschner (1988c) listed this species only from Manitoba and Montana from our region, and Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. This species has been recorded from a variety of different plant species; my survey works agrees with this as most specimens were collected in Old Field situations with quite a diversity of weed species. The life history of this species has been studied (Oetting and Yonke 1971b), and the immature stages have been described (Es- selbaugh 1946, DeCoursey and Esselbaugh 1962). Specimens Examined: MINNESOTA [New State Record]: Clay Co.: Bluestem Prairie, 46°50′Lat., 96°26′Long., 10-IX-1999, C. Davis, C. Jordan (1♀ NDSU); Bluestem Prairie, 4mi. E, 1 mi. S Glyndon, 46°50.08′N, 96°25.08′W, 24-IX-1999, P. B. Beauzay (1♂ NDSU); Regional Science Center Buffalo River Site, 29-VIII-2002, F. Antwi (1♀ NDSU). Marshall Co.: 4mi. W Argyle, 27- IX-2008, K. Rivard (1♂ NDSU). NORTH DAKOTA: Barnes Co.: S of Valley City, 47°04.00′N, 98°04.00′W, 18-IX-2002, A. Joshi (1♀ NDSU). Burke Co.: 17- VIII-1958, S. Wilson (1♀ NDSU). Burleigh Co.: Double Ditch, 27-IX-1975 (1♀ UNDG). Cass Co.: Fargo, 17-IX-2008, D. Hanson (1♀ NDSU), 18-IX-1921, R. L. Webster (1♀ NDSU), 21-IX-1958, G. J. & R. L. Post (12♂♂ 19♀♀ NDSU), Fargo, NDSU campus, 21-V-2012, D. A. Rider (3♀♀ DARC), 25-VI-2012 (1♂ DARC), 24-VIII-2002, F. Antwi (1♀ NDSU), 3-IX-2008, D. A. Rider (1♂ 1♀ DARC), 6-IX- 2001, D. A. Rider (1♂ 2♀♀ DARC), 11-IX-2008 (1♀ NDSU), 17-IX-2003, C. J. Kjergaard (1♀ NDSU), 25-IX-2000, B. Gebhard (1♂ NDSU), 9-X-2003, B. Foster (1♂ NDSU); Wild Rice R. at Red Riv., 46.758N, 96.787W, 1-VIII-2012, D. Rider (1♀ DARC); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (1♀ DARC), 1-VIII-2012, D. A. Rider (3♂♂ 3♀♀ DARC), 13-VIII-2012, D. A. Rider (3♂♂ 2♀♀ DARC); 27-V-2002, J. Knott (1♀ DARC); 1958 (1♂ NDSU). Cavalier Co.: 3-VIII-1961, R. L. Post (1♀ NDSU). Dunn Co.: 13-VII-1976, L. A. Schutz & J. W. Smith (1♀ NDSU), 28-VII-1976, Lago & Kurtz (1♀ NDSU). Grand Forks Co.: Air Force Base Property, McKinock Twp, T152N, R53W, Sec. 27, 16-IX-1970, D. Nelson (3♂♂ 1♀ UNDG); Arvilla, 28-IX-1931 (1♂ 2♀♀ UNDG), X-1929 (1♂ UNDG); Forest River Biol. Area, 1.8mi. W, 1mi. N Inkster, T154N, R55W, Sec. 11 & 14, 18-IX-1976 (1♂ UNDG), 5-X-1969 (1♀ UNDG); Grand Forks, 30-IX-1963 (1♀ UNDG), 16-X-1931 (1♀ UNDG), 16-X-1950 (1♀ UNDG); Northwood, 17-VI-1959, R. L. Post (1♂ NDSU); Oakville Prairie, Oakville Twp, Sec. 9, 2-X-1959 (1♀ UNDG); Oakville Prairie, T151N, R52W, Sec. 16, 26-IX-1983 2012 THE GREAT LAKES ENTOMOLOGIST 345 (1♂ UNDG); Thompson, 1.5mi. W of I-29 interchange, 20-IX-1992, J. Froelich (2♀♀ UNDG); Turtle River State Park, 30-VI-1953 (1♂ UNDG), 2-X-1992 (1♀ UNDG). Hettinger Co.: 19-VIII-1977, L. A. Schutz & J. W. Smith (1♀ NDSU). McKenzie Co.: Th. Roos. M. Prk, 5-VI-1965, L. Grochowski (1♂ NDSU), 23-VI- 1965, L. Grochowski (1♀ NDSU). Mercer Co.: Beulah, 3-IX-1977 (1♀ UNDG); 18mi. S, 4mi. W Beulah, 3-IX-1977 (1♀ UNDG); Stanton area, 1-VII-1972, A. C. F. Hung (2♂♂ 1♀ UNDG), 4-6-VIII-1972, A. C. F. Hung (2♂♂ 3♀♀ UNDG); T144N, R84W, Sec. 22, 1-VII-1972, A. C. F. Hung (1♂ UNDG), 15-VIII-1972, P. B. Kannowski (1♂ UNDG); T144N, R85W, Sec. 9, 18-20-VII-1972, A. C. F. Hung (1♀ UNDG). Mountrail Co.: Clearwater Lake, 17-IX-1998, A. Nelson (1♀ MISU); 19-VIII-1958, S. Wilson (3♀♀ NDSU). Morton Co.: Mandan, VII-1965, J. W. Goertel (1♀ NDSU). Oliver Co.: T143N, R83W, Sec. 6, 16-VIII-1972, P. B. Kannowski (1♂ UNDG). Pembina Co.: 8-VIII-1961, R. L. Post (1♀ NDSU), 11-VIII-1958, S. Wilson (1♀ NDSU), 11-IX-1958, S. Wilson (1♀ NDSU). Ransom Co.: Mirror Pool WMA, 25-IX-1993, M. J. Dietrich (1♀ NDSU). Renville Co.: 15-VIII-1958, S. Wilson (1♀ NDSU). Richland Co.: 1mi. SE McLeod, 26-V-1992, J. R. Powers (1♀ DARC); Mirror Pool, 23-VIII-1974, P. K. Lago (1♀ NDSU); 16- V-1962, R. Frye, alfalfa (1♀ NDSU), 21-IX-1957, R. L. Post & M. Olson (1♂ 4♀♀ NDSU); SNG, Oak Savannah, T135N, R52W, Sec. 10, 7-VIII-1993, D. Cuthrell (1♀ NDSU). Sargent Co.: Cayuga, 19-VI-1962, R. D. Frye (1♀ NDSU). Slope Co.: Burning Coal Vein St. Pk., 19-VII-1978, E. U. Balsbaugh (1♀ NDSU). Stark Co.: Patterson Lake Dickinson, 14-VI-2011, D. Nelson, Lindgren trap (1♀ NDSU). Traill Co.: 10-VI-1958 (1♀ NDSU). Ward Co.: Minot, 26-VI-1936, M. Rita (1♀ MISU); SE Minot, behind Menards, 17-IX-1999, G. Hanley (1♀ MISU). Wells Co.: Harvey, 3-VII-1961, B. Liebe (1♂ MISU). Holcostethus macdonaldi Rider and Rolston, 1995 - This is the only species in North Dakota with the abdominal venter dark brown with a few pale markings; all other species have the abdominal venter pale, sometimes with a few dark markings. Until recently, this species went under the name H. piceus, but this name is preoccupied; Rider and Rolston (1995) provided the replace- ment name macdonaldi. McPherson (1982, as H. piceus) gave the distribution as Québec west to Alberta, and south to Illinois and Colorado. This species has not yet been recorded from North Dakota, although Froeschner (1988c) and Maw et al. (2000) reported it from all surrounding states or provinces except Minnesota. This species appears to be much scarcer than the other two species reported from North Dakota. Specimens Examined: NORTH DAKOTA [New State Record]: Barnes Co.: Valley City St. Univ., 17-X-1976 (1♂ UNDG). Benson Co.: T156N, R71W, Sec. 10, 2-V-1962, P. E. Slabaugh (1♂ NDSU). Burleigh Co.: Bismarck landfill site, 29-VII-2008, D. Nelson (3♀♀ NDSU). Cass Co.: VII-1957 (1♂ NDSU). Mercer Co.: T144N, R84W, Sec. 28, 18-20-VII-1972, A. C. F. Hung (1♀ UNDG); T144N, R84W, Sec. 22, P. B. Kannowski, 8-VIII-1973 (1♂ UNDG). Morton Co.: Ft. Abraham Lincoln St. Pk., 28-VII-2007, D. Nelson, Lindgren trap (1♀ NDSU). Richland Co.: SNG, Horsetrail Head nr. Oak Savannah, 25-X-2007, D. A. Rider (1♀ DARC).

Menecles Stål, 1867 This genus contains only two species, both North American. Menecles portacrus Rolston is only known from the Big Bend area of Texas; M. insertus (Say) is more widespread and now known to occur in North Dakota. Rolston (1972b) reviewed the genus. Menecles insertus (Say, 1832) - This species is fairly easy to recognize simply by the body shape; it is more ovoid than most pentatomids, somewhat flattened, and the anterolateral pronotal margins are distinctly convex. The distribution of this species was given by McPherson (1982) as Québec, Ontario, 346 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 and New England west to Nebraska and Kansas, southwest to Arizona and Cali- fornia, and south to Arkansas and Florida. This species has not been recorded from North Dakota or Minnesota. The closest records given by Froeschner (1988c) are from Illinois and Nebraska. The life history has been studied by Balduf (1945), and the immature stages have been described (Esselbaugh 1946, DeCoursey and Esselbaugh 1962). Specimens Examined: MINNESOTA [New State Record]: Becker Co.: 6mi. SW Detroit Lakes, 24-29-VII-1983, M. Moher (1♂ DARC). Clay Co.: Moor- head, 27-IX-1989, J. R. Powers (2♂♂ DARC). NORTH DAKOTA [New State Record]: Grand Forks Co.: Turtle River, 4.8mi. W, 2mi. N Emerado, T152N, R53W, Sec. 20 & 29, 24-IX-1976 (1♂ UNDG). McKenzie Co.: 25-V-1961, R. L. Post (1♀ NDSU). Stark Co.: Dickinson, Patterson Lake, 13-VII-2011, K. Marmon, Lindgren trap (1♀ NDSU).

Mormidea Amyot and Serville, 1843 This is another large, diverse New World genus with many species distributed primarily throughout the neotropics. Other than a few species that enter into the United States in extreme southern Florida or the desert southwest, M. lugens (Fabricius) is the only widespread species in North America. Rolston (1978) revised the genus Mormidea. Mormidea lugens (Fabricius, 1775) - Mormidea lugens is easy to separate from other North Dakota pentatomoids by virtue of being the only smallish, dark brown species with a distinct ivory callus transversing the anterior disk of the pronotum. McPherson (1982) gave the distribution of this species as On- tario, Québec, and Maine south to Florida and west to North and South Dakota, Wyoming, Colorado, Oklahoma, and Texas. The only literature record from the Upper Midwest I have found is that of Uhler (1876), who recorded this species from “Dakota” and “Indian Terr.” Froeschner (1988c) listed this species from North Dakota, perhaps based upon the above “Dakota” record. Maw et al. (2000) also recorded this species from Manitoba. The records presented herein appear to be the first specific literature records for either North Dakota or Minnesota. The records presented would seem to indicate that this species occurs across most of Minnesota at least as far north as the Itasca area (Clearwater Co.); in North Dakota, the known records are confined to the southeastern corner of the state. The immature stages have been described (Esselbaugh 1946, DeCoursey and Esselbaugh 1962). McPherson (1974) studied the biology of this species. Specimens Examined: MINNESOTA [New State Record]: Clearwater Co.: Itasca State Park, 24-VI-1958, R. Gundersen (1♀ DARC). Fillmore Co.:24- VI-1984, M. Tenney (1♀ DARC). Hubbard Co.: Camp Wilderness, 23-VII-1995, G. Fauske (1♂ NDSU). Morrison Co.: 10mi. N Little Falls, 17-VI-1981 (1♀ DARC). Norman Co.: Agassiz Dunes SNA, 47°29’Lat. 96°18’Long., 27-V-1999, P. Tinerella, G. Fauske, pitfall (1♀ NDSU), 11-VI-1999, P. Tinerella, G. Fauske, pitfall (2♀♀ NDSU), 8-VII-1998, P. Tinerella, pitfall (1♀ NDSU), 17-VII-1996, K. Urlacher, C. Locken, pitfall (2♀♀ NDSU); Agassiz Dunes SNA, 0.5mi. W, 2mi. S Fertile, 47°30.02’N, 96°18.36’W, 27-V-1999, Davis, Jordan (1♂ NDSU); 2mi. S Fertile, 47°29.92’N, 96°18.57’W, 28-VI-2001, P. Beauzay (1♂ NDSU). Polk Co.: Agassiz Dunes SNA, 47°30’Lat. 96°18’Long., 1-VII-1997, P. Tinerella, A. Abbott, pitfall (3♀♀ NDSU), 8-VII-1998, P. Tinerella, pitfall (1♀ NDSU), 9-VII-1999, P. Tinerella, G. Fauske, pitfall (2♀♀ NDSU), 21-VII-1997, A. Abbott, P. Tinerella, pitfall (6♀♀ NDSU); 31-VII-1998, P. Tinerella, pitfall (2♀♀ NDSU). NORTH DAKOTA [New State Record]: Cass Co.: Prosper NDSU Res. Plots, T141N, R51W, Sec. 27, 3-VII-2002, L. M. Tobias (1♂ NDSU); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (1♀ DARC). Ransom Co.: Mir- ror Pool WMA, 46°32.348N, 97°17.475W, 27-VI-2012, D. A. Rider (1♂ DARC). Richland Co.: Ekre’s Grassland Preserve, 46.552N, 97.130W, 28-V-2010, D. 2012 THE GREAT LAKES ENTOMOLOGIST 347 A. Rider (1♀ DARC); 28-VII-1993, D. L. Cuthrell (1♀ NDSU); 0.5mi. SE Mirror Pool, 6-VI-1995, D. A. Rider (3♀♀ DARC); 1mi. E Mirror Pool, 28-VII-1993, D. A. Rider (1♀ DARC); 1mi. NE Mirror Pool, 17-VII-1993, D. L. Cuthrell (2♂♂ NDSU); 1.5mi. NE Mirror Pool, 15-VI-1994, D. A. Rider (2♂♂ 3♀♀ DARC); 2mi. NE Mirror Pool, 28-VI-1993, G. Fauske (1♂ 2♀♀ NDSU); SNG, Oak Savannah area, T135N, R52W, Sec. 10, 6-VI-1995, C. Locken, D. Pitzen, G. Fauske (1♂ NDS), 11-VI-1996, L. DeCock (3♂♂ 6♀♀ NDSU), 13-VI-1995, G. Fauske, C. Locken, D. Pitzen (1♂ 2♀♀ NDSU), 18-VI-2000, D. A. Rider (1♂ 1♀ DARC), 17-VII-1993, D. Cuthrell (2♀♀ NDSU); SNG, Horsetrail Head, 46°31.409N, 97°12.194W, 27-VI-2012, D. A. Rider (1♂ 2♀♀ DARC), 25-VIII-2009, D. A. Rider (3♂♂ DARC), 4-IX-2009, D. A. Rider (4♂♂ 3♀♀ DARC); SNG, Horsetrail Head, 46.523N, 97.201W, 28-V-2010, D. A. Rider (2♂♂ 7♀♀ DARC); SNG, Horse trail head, 46.52N, 96.21W, 20-VII-2012, D. A. Rider (1♂ 2♀♀); T136N, 61W, Sec. 32, 23-VII-1993, G. Fauske (2♂♂ 1♀ NDSU).

Trichopepla Stål, 1867 Members of this genus are easily recognized by the pubescent dorsum. There are other genera with species that are pubescent, but none of these occur in North Dakota. McDonald (1976) recognized seven species (all of which occur in North America) in his revision of Trichopepla. Only one species, T. atricornis Stål, is known to reach the Upper Great Plains. The widespread T. semivittata (Say) does not occur as far north as North Dakota, the closest known records coming from Iowa and Nebraska. The unlocalized record of Trichopepla grossa Van Duzee from South Dakota (McDonald 1976) needs confirmation; I have not been able to locate the source of this record. The remaining species are all distributed in the western and/or southern United States. In the event that T. semivittata and/or T. grossa might actually occur in the southern parts of North Dakota, a key is provided for the separation of these three species.

Key to species of Trichopepla that may be found in North Dakota 1 Connexiva pale with dark mesial margins, the dark markings also along both anterior and posterior connexival incisures, producing a distinct alternating pale and dark pattern; if connexival pattern is weak, then second antennal segment is pale, and head is somewhat elongate, tapering to a narrowly rounded apex [not yet found in North Dakota]...... semivittata (Say) – Connexiva pale with dark mesial margins, the dark markings sometimes slightly intruding along connexival incisures, at most creating an undulat- ing pattern; second antennal segment black, somtimes pale at joints; head shorter, not tapering apically, broadly rounded apically...... 2 2(1) Anterolateral pronotal margins reflexed to narrowly explanate, distinctly incised submarginally...... atricornis Stål – Anterolateral pronotal margins edged, but not reflexed or explanate, not incised submarginally [not yet found in North Dakota]...... grossa Van Duzee Trichopepla atricornis Stål, 1872 - Trichopepla atricornis is easy to separate from all other North Dakotan pentatomids by the densely pubescent dorsal surface of the body. McPherson (1982) gave the distribution as Ohio, Illinois, and Wisconsin west to Alaska, British Columbia, and California. This species has not officially been recorded from either Minnesota or North Dakota, even though Froeschner (1988c) listed it from Montana and several states both to the east and west of North Dakota; also Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. Members of Trichopepla seem to show a preference for plants in the Apiaceae. 348 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Specimens Examined: MINNESOTA [New State Record]: Clay Co.: Bicentennial Prairie, 47°03′Lat., 96°25′Long., 9-VII-1997, P. Tinerella, A. Ab- bott, pitfall (1♀ NDSU), 28-VII-1997, P. Tinerella, pitfall (1♂ NDSU); Blazing Star Prairie, 47°03′Lat., 96°25′Long., 1-V-1999, P. Tinerella, G. Fauske, pitfall (1♂ NDSU), 27-VII-1998, P. Tinerella (1♀ NDSU); Bluestem Prairie, T139N, R45W, Sec. 19, 3-IX-1996, J. Albertson, L. DeCock (1♂ 2♀♀ NDSU); Bluestem Prairie, T139N, R46W, Sec. 10, 3-IX-1996, J. Albertson, L. DeCock (1♀ NDSU), Sec. 15, 13-VII-1995, D. Rider, C. Locken, G. Fauzke (4♂♂ 6♀♀NDSU), Sec. 21- 15, 17-VII-1995, D. Rider, C. Locken, G. Fauske (1♂ NDSU); Bluestem Prairie, 46°50′Lat., 96°25′Long., 1-X-1998, P. Tinerella, pitfall (1♀ NDSU); Bluestem Prairie, 46°51′Lat., 96°25′Long., 31-VII-1997, P. Tinerella, A. Abbott, pitfall (1♀ NDSU); western edge Bluestem Prairie, 10-IX-2009, D. A. Rider (9♂♂ 11♀♀ DARC); B-B ranch, 47°02′Lat., 96°26′Long., 9-VII-1997, P. Tinerella, A. Abbott, pitfall (1♀ NDSU). NORTH DAKOTA [New State Record]: Grand Forks Co.: Oakville Prairie, Oakville Twp, Sec. 9, 29-VI-1966 (1♂ UNDG). Mercer Co.: T144N, R85W, Sec. 9, 5-VI-1973, P. B. Kannowski (1♂ UNDG). Oliver Co.: T143N, R83W, Sec. 6, 15-VIII-1972, P. B. Kannowski (3♂♂ UNDG). Williams Co.: 1mi. N Williston, 1-X-1977 (1♀ UNDG).

Tribe Halyini Amyot and Serville, 1843 The tribe Halyini is mostly an Old World tribe with only two genera known from the New World; it is especially diverse in Africa, the Orient, and Austra- lia. Defining characters, other than those associated with the male genitalia, are few and difficult to discern. Gross (1976) gives a fairly detailed discussion of the genital characters. It should also be noted that at one time many New World genera from Central and South America were included in the Halyini, but all of them except the two treated in this paper have now been transferred to the tribe Ochlerini in the subfamily Discocephalinae. The two remaining genera, Brochymena and Parabrochymena, are both mainly Nearctic genera with a handful of species in Mexico, and a couple more reaching Costa Rica and Panama. They generally are very cryptic, colored so as to blend in with the bark of various tree species.

Key to the genera of Halyini occurring in North Dakota 1 Humeral angles subquadrate; basal disk of scutellum distinctly elevated...... Parabrochymena – Humeral angles subtriangular, rounded to angulate, but not quadrate; basal disk of scutellum at most slightly elevated...... Brochymena

Brochymena Amyot and Serville, 1843 This genus, and the recently split-off genus Parabrochymena, are the only New World representatives of the tribe Halyini. Their species have a somewhat flattened appearance, especially ventrally, and are usually mottled black, gray, and white. It is easy to visualize these species blending onto the bark on various tree species, where they are most commonly collected. Brochymena was recently divided into two genera; the species having quadrate humeral angles were transferred to Parabrochymena (Larivière 1992). Acceptance of this division has not been universal, but it seems best to follow this classification until a more thorough study has been completed (see McPherson and Ahmad 2005, 2007, for further discussion). Brochymena contains a number of North American species, several of which also occur in Mexico and into Central America. Only one species reaches into North Dakota however. Herbert Ruckes published a series of papers (1938, 1939a, b, c, 1941, 1946) describing several new species, and the biology or feeding habits, finally culminating in a revision (1947) of the 2012 THE GREAT LAKES ENTOMOLOGIST 349 genus. Even though Ruckes’ paper was extremely detailed, he designed identification keys that also reflected phylogeny, making them incredibly difficult to use. More recently, Larivière (1992) published an excellent review of the genus. Brochymena quadripustulata (Fabricius, 1775) - This is the only species of Brochymena in North Dakota, so it is fairly easy to recognize by the generic characters. It and Parabrochymena arborea both have the tooth near the apex of each jugum, but this species has the subtriangular humeral angles (subquadrate in P. arborea). It also has the juga much longer than the tylus, and usually meeting anterior to the tylus. There are other species of Brochy- mena that are difficult to separate fromB. quadripustulata in other areas of the U.S.; in fact, examination of the male genitalia is necessary for some species. The distribution given by McPherson (1982) is across southern Canada and the continental United States, from the Atlantic to Pacific coasts, and ranges south into northern Mexico. Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan; Froeschner (1988c) did not list this species from North Dakota, or from any of the surrounding states or provinces, even though Lugger (1900, as B. annulata) recorded this species as a pest of apple trees in Minnesota. Larivière (1992) listed B. quadripustulata from both Minnesota and North Dakota, but did not give any specific localities. Cuda and McPherson (1976) studied the life history of this species, including rearing it in the laboratory, and they provided a description of the immature stages. Gamboa and Alcock (1973) studied the mating behavor. Specimens Examined: MINNESOTA: Clay Co.: Bluestem Prairie, western edge, 10-IX-2009, D. A. Rider (1♀ DARC); Moorhead, 27-IX-1989, J. R. Powers (1♂ DARC); Moorhead Science Center, 26-VIII-2004, S. Ganehiarach- chi (1♀ NDSU); 4-IX-2008, D. A. Rider (3♂♂ 1♀), 6-IX-2007, D. A. Rider (2♂♂ 1♀). Stearns Co.: St. Cloud, 18-IX-1983, Garder (1♂ DARC), 18-IX-1984, M. Nelson (1♀ DARD), 29-IX-1982, Y. Ito (1♀ NDSU); St. Cloud, State College, 15- VII-1967 (1♀ NDSU), 2-X-1968 (1♂ DARC). NORTH DAKOTA: Billings Co.: vicinity Medora, 21-VI-2000, G. Hanley (1♀ MISU). Burleigh Co.: Bismarck, 1-V-1987, R. Wilmont (1♀ NDSU); 19-IX-1985, D. Cashman (1♂ NDSU). Cass Co.: Fargo, 8-VI-1923, W. Baker (1♂ NDSU). Divide Co.: 11-IX-1962, R. L. Post & R. D. Gordon, beating dead box elder (1♂ NDSU). Dunn Co.: 25-VIII- 1976 (1♀ NDSU). Grand Forks Co.: Grand Forks, 2-X-1975 (1♂ UNDG); State Park near Arvilla, 1-X-1969 (1♂ UNDG); Turtle R., 4.8mi. W, 2mi. N Emerado, T152N, R53W, Sec. 20 & 29, 7-IX-1977 (1♂ UNDG), 8-IX-1982, K. Kreil (1♂ 1♀ UNDG); UNDG Biology area Forest R. Valley, Inkster, 3-X-1965 (1♀ UNDG). Kidder Co.: Lake Isabel, 20-VIII-1980 (1♂ UNDG). McHenry Co.: Sec. 36, T156, R78, 26-V-1967, P. E. Slabaugh (1♂ NDSU). McIntosh Co.: 22-IX-1962, R. L. Post & G. Thomasson (1♀ NDSU). McLean Co.: Ruso, 3-VII-1938, E. Boe (1♂ MISU). Morton Co.: Mandan, 9-IX-1985, T. Schmidt (1♀ NDSU), 22-IX-1922, R. L. Webster (1♂ NDSU). Mountrail Co.: Sanish, 16-V-1973 (1♀ NDSU). Oliver Co.: Sec. 6, T143N, R83W, 16-VIII-1972, P. B. Kannowski (1♀ UNDG). Pembina Co.: 6-VII-1961, E. Saugstad (1♀ NDSU). Ransom Co.: Lisbon, summer 1983, J. Julian, on leafy spurge (1♀ NDSU); SNG, A Annex site, T136N, R52W, NW8, 17-VII-1992, D. A. Rider (1♀ DARC). Renville Co.: 11-IX-1962, R. L. Post & G. L. Thomasson, beating dead thornapple (1♂ NDSU). Richland Co.: Ekre Grassland Pres., 28-VIII-2008, D. A. Rider (1♀ DARC); Hankinson, 22-IX-1923, R. L. Webster (1♂ NDSU); Mirror Pool, 4-IX-1998, D. A. Rider (1♀ DARC). Slope Co.: Burning Coal Vein, 7-VI-1973, T. L. McCabe (1♂ NDSU). Stark Co.: Dickinson, Patterson Lake, 14-VI-2011, D. Nelson, Lindgren trap (1♂ NDSU), 28-VI-2011, K. Marmon, Lindgren trap (1♂ NDSU), 13-VII-2011, K. Marmon, Lindgren trap (2♂♂ 1♀ NDSU), 26-VII-2011, K. Marmon, Lind- gren trap (1♂ NDSU), 23-VIII-2011, K. Marmon, Lindgren trap (2♀♀ NDSU). Ward Co.: Minot, 25-IV-1936, E. Schrel (1♀ MISU), 17-V-1939, L. Hanson (1♀ MISU), 10-VII-1936, S. Clotidis (1♀ MISU), 13-VII-1993, R. Harry (1♀ MISU); NW Minot, 13-VI-2003, G. Hanley, sweeping alflafa (1♂ MISU); 17-XI-1962, R. 350 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

L. Post & R. Gordon (1♂ NDSU). Williams Co.: Williston, 1mi. N, 2-VI-1978 (1♂ UNDG), 3-IX-1977 (1♂ 1♀ UNDG).

Parabrochymena Larivière, 1992 This genus is very closely related, and may actually be congeneric with Brochymena. Both are somewhat flattened gray to fuscous species mottled with pale markings so as to easily blend in with tree bark. There are apparently genitalic differences, but the easiest way to separate the two genera is by the form of the humeral angles. Parabrochymena species tend to have subquadrate humeral angles whereas they are more angular to rounded in Brochymena. This genus was revised by Larivière (1994). There are several southern and western species of Parabrochymena, but only a single species extends as far north as North Dakota. Parabrochymena arborea (Say, 1825) - Similar to B. quadripustulata, this is the only species of Parabrochymena to occur in North Dakota, so identification is fairly easy by using generic characters. The juga with a tooth near the apex, coupled with subquadrate humeral angles should separate this species from all others in North Dakota. McPherson (1982, as Brochymena arborea) provided the following range: Québec, Ontario, and New England south to Florida, and west to Kansas, New Mexico, Mexico, South Dakota, and Iowa. Neither Uhler nor Hussey recorded this species from North Dakota; Froeschner (1988c) listed it from South Dakota. Larivière (1994) provided a list of states (without specific localities) in which this species had been collected; her list included Minnesota and South Dakota, but not North Dakota. The distribution map she provided, however, had the southern half of North Dakota shaded. I consider the following record to be the first official record for North Dakota. To date, only a single specimen is known from North Dakota. I have also provided a couple records from Minnesota because its northern distribution may not be well defined. Specimens Examined: MINNESOTA: Becker Co.: Detroit Lakes, 2-IX- 1984, V. Beregovoy (1♀ NDSU). Morrison Co.: 10mi. N Little Falls, 21-VII-1981 (1♀ NDSU), 1-VIII-1981 (1♀ DARC). Ottertail Co.: Maplewood State Park, 29-IX-2001, L. M. Tobias (1♂ NDSU). Stearns Co.: St. Cloud, 1-X-1984, M. Nelson (1♂ DARC); St. Cloud, State College, 15-X-1965 (1♀ NDSU). NORTH DAKOTA [New State Record]: Cass Co.:Absaraka Arboretum, 14-IX-2000, A. Hackey (1♀ NDSU).

Tribe Mecideini Distant, 1902 The Mecideini includes a single genus, Mecidea, but has species occurring throughout the Old and New Worlds. It is characterized primarily by the stridulatory areas on the abdominal venter, although such areas are also present in members of the Diemeniini, a primarily Australian tribe. The mecideines are elongate, slender species, feeding primarily on grasses in somewhat arid areas; the diemenines are more typically stink-bug shaped, and not definitely associated with grasses.

Mecidea Dallas, 1851 This genus has species in both the New World and Old World; they tend to occur on grasses in xerophytic regions. In the New World, there are four known species: M. longula occurs in the West Indies, M. pampeana occurs in southern South America, and M. major and A. minor both occur in southwestern and central states of the U.S., with M. minor occurring as far south as northern South America. The American species were once confused with M. longula 2012 THE GREAT LAKES ENTOMOLOGIST 351 and much of the earlier literature for both species is under that name. Sailer (1952) revised the genus. Species of this genus are easily recognized by the pale yellowish color, and by the presence of stridulatory ridges on the abdominal venter; the only other North American genus that is elongate, slender, and pale yellow is Oebalus, but the widespread O. pugnax (which lacks the stridulatory areas, and has spinose humeral angles) does not occur in our area. No other North American pentatomid genera have stridulatory ridges on the abdominal venter. Mecidea major and M. minor can be difficult to separate, but M. major does not occur in our area (nearest records are from Illinois and Missouri). Just in case, however, I have provided some distinguishing characters. In M. minor, the spiracles are usually pale, there may be a row of dark markings along the midline of the abdominal venter, the males have a tubercle near the posterior margin of the pygophore, and the females have a relatively small black spot on the abdominal venter just below each abdominal setigerous puncture. In M. major, the spiracles are often brown to black, the row of black spots along the midline of the abdominal venter are usually lacking, the males lack the tubercle near posterior margin of pygophore, and the females have a relatively large black spot on the abdominal venter just below each setigerous puncture. Mecidea minor Ruckes, 1946 - This species is very easy to recognize; it is the only elongate, slender, pale yellowish pentatomid known to occur in North Dakota. McPherson (1982) gave the range of this species as northern Mexico north to central California, northern Utah, and central South Dakota, with the eastern boundary near the eastern borders of Texas, Oklahoma, and Kansas. This species has not previously been recorded from North Dakota. Harris (1943) recorded this species (as M. longula) from central South Dakota. At present, only a single specimen is known from North Dakota. Specimen Examined: NORTH DAKOTA [New State Record]: Cass Co.: Wild River at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (1♀ DARC).

Tribe Nezarini Atkinson, 1888 It is very risky to make determinations based on color characters; it is even more risky to base classifications on color characters. That is why I am hesitant to say that the easiest way to recognize the tribe Nezarini is that its members are green in color, and usually remain green after death. There are other tribes whose members are green while alive (some Procleticini, Rhynchocorini, and a undescribed tribe related to Chlorocoris), but they invariably turn pale yellow after death. As presently understood, this tribe contains a number of genera, both in the Old World and New World. Several of these occur in the United States; I have treated four North Dakota genera in this tribe, but I have some concerns about including Tepa and Thyanta.

Key to the Genera of Nezarini occurring in North Dakota 1 Base of abdominal venter produced forward into a small obtuse spine or tubercle...... Chinavia – Base of abdominal venter smoothly rounded, not produced forward..2 2(1) Smaller, usually less than 8mm in length...... Tepa – Larger, usually more than 10mm in length...... 3 3(2) Ostiolar ruga elongate, reaching at least two-thirds to edge of supporting plate, and acuminate apically...... Thyanta – Ostiolar ruga much shorter, reaching no more than half way to edge of supporting plate, not acuminate apically...... Chlorochroa 352 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Chinavia Orian, 1965 This genus has had a somewhat checkered taxonomic history, and has only recently become accepted. Most of the species included in this genus were considered to be members of the genus Acrosternum. It now appears that the true Acrosternum should be confined to those Old World species that are somewhat smaller in size, paler in color, and occur in more arid regions. In contrast, species of Chinavia are larger in size, much brighter green in color, and they occur in temperate to tropical areas in both the New World and Africa. This is a very diverse genus with many North American and Neotropical species. The genus was recently revised (Rolston 1983), and there have been a number of recent additions, none of which affects North American species. There are two species which are relatively widespread in eastern North America, but one, C. pennsylvanica (DeGeer), does not occur in North Dakota (it is much more broadly rounded, with distinctly convex anterolateral pronotal margins); the closest records are from Iowa and Missouri (there is actually a record of this species from Minnesota, but this is based upon a misidentification - see below). Chinavia hilaris (Say, 1832) - This species, commonly referred to as the green stink bug, is easily recognized by its somewhat larger size, the long, acuminate ostiolar ruga, and the basal, anteriorly directed spine or tubercle on the abdominal venter. The distribution given by McPherson (1982) was from Québec and New England west through southern Canada and the northern states to the Pacific coast, and south and southwest to Florida, Texas, Arizona, Utah, and California. Froeschner (1988c) listed this species from Montana and South Dakota. Lugger (1900), in Minnesota, briefly described the biology of what he called Acrosternum pennsylvanicum. It is, however, evident from his brief description and the accompanying figure that he actually had specimens of this species. It appears that this common species has not been recorded from North Dakota. All North Dakota records are from the southeastern part of the state except for the Nelson Co. record which is a little farther north. Because the northern limits of this species are poorly known, I have also provided several collection records from Minnesota. McPherson and Tecic (1997) studied the life history of this species. Specimens Examined: MINNESOTA: Becker Co.: Detroit Lakes, 26-VIII-1984, V. Beregovoy (1♂ NDSU); 6mi SW Detroit Lakes, 23-V-1990, P. Kamrud (1♂ DARC), 5-VI-1990, J. R. Powers (1♀ DARC). Big Stone Co.: Clinton, 10-IX-2002, N. Brandt (1♂ NDSU). Clay Co.: Moorhead Science Center, 6-IX-2007, D. A. Rider (2♂♂ 1♀ DARC). Ottertail Co.: 12-VI-2008, K. Johnson (1♂ UNDG). Scott Co.: UM Landscape Arboritum, 27-IX-2008, J. Ell (1♂ NDSU). Stearns Co.: Cold Spring, 1-X-1972, W. Karulak (1♀ DARC); State College, St. Cloud, 25-VII-1965 (1♀ DARC). NORTH DAKOTA [New State Record]: Cass Co.: Absaraka Arboretum, 14-IX-2000, A. Hackey (1♀ NDSU); Fargo, NDSU campus, 2-IX-2008, D. A. Rider (1♂ DARC). Grand Forks Co.: 12-IX-2008, C. Black (1♂ UNDG), 18-IX-2008, C. Hanson (1♂ UNDG). Nelson Co.: 2-VII-1958 (1♀ NDSU). Pembina Co.: Crystal, 12-VIII- 1989, M. Dietrich (1♀ NDSU). Ransom Co.: SNG, A-Annex, 19-V-2005, G. M. Fauske, Merc. vapor/blklight (1♀ DARC); Mirror Pool WMA, 30-VIII-2006, C. P. Johnson (1♂ NDSU), 30-VIII-2007, D. A. Rider (1♂ 2♀♀ DARC); SNG, Sagvold, 23-VII-1993, D. Cuthrell, Hg vapor light (1♂ DARC). Richland Co.: Jerome Link Farm, 21-X-2002, T. Link (1♀ NDSU); Mirror Pool, 6-VI-1993, J. Nelson, blk light (1♂ NDSU), 4-IX-1998, D. A. Rider (1♀ DARC), 11-IX-1974, R. K. Lago (2♂♂ NDSU). Chlorochroa Stål, 1872 Chlorochroa species tend to be medium in size (smaller than Chinavia, but larger than Tepa or Thyanta), the ostiolar ruga is short and auriculate, and the abdominal venter lacks the basal spine or tubercle. Several species within 2012 THE GREAT LAKES ENTOMOLOGIST 353 this genus are very difficult to separate. Fortunately, there have been three relatively recent publications, each with many illustrations and identification keys (Buxton et al., 1983; Thomas, 1983; Scudder and Thomas, 1987).

Key to North Dakota species of Chlorochroa 1 Scutellum with more or less distinct medial yellow longitudinal stripe; scutellar tongue marked with brown or black on each side...... 2 – Scutellum without medial yellow longitudinal stripe; scutellar tongue not marked with brown or black...... 3 2(1) Relatively small, less than 12mm in length; outer margin of each embolium lacking dark brown or black markings...... Chlorochroa faceta (Say) – Relatively large, more than 12mm in length; outer margin of each embolium narrowly margined with brown or black...... Chlorochroa belfragii (Stål) 3(1) Relatively small, length less than 10mm...... Chlorochroa viridicata (Walker) – Relatively large, length greater than 11mm...... 4 4(3) Base of scutellum with three more or less distinct pale spots, one in each basal angle and one medial; embolium only slightly wider apically than basally; angle between posterior and medial processes of each paramere much greater than 90 degrees...... Chlorochroa uhleri (Stål) – Basal spots of scutellum lacking or very small; embolium much wider apically than basally; angle between posterior and medial processes of each paramere about 90 degrees...... 5 5(4) Male pygophore in posteroventral view with small central tumescence...... Chlorochroa ligata (Say) – Male pygophore in posteroventral view with well-developed subapical transversely arcuate ridge, lacking central tumescence...... Chlorochroa persimilis Horváth Chlorochroa belfragii (Stål, 1872) - This species is fairly easy to recognize. There are only two pentatomid species in North Dakota that are green to gray-green with a yellowish, longitudinal stripe down the dorsal midline. The other species is C. faceta, which tends to be smaller than C. belfragii. Also, C. faceta lacks the dark markings on the lateral margins of the embolium that are characteristic of C. belfragii. This species is relatively widespread, but is rarely collected; in fact, at one time it was a candidate species for the U.S. threatened or endangered lists. It may not be as rare as previously thought; it may just be under collected because of its biology. This species belongs in the subgenus Rhytidolomia, and is related to C. dismalia Thomas, C. saucia (Say), and C. senilis (Say). Both C. saucia and C. senilis occur on salt marsh grasses along the Atlantic and Gulf coasts, and Parshley (1923) reported findingC. saucia down in the matted grasses near the base of the plants. The specimens I collected on Oakville Prairie were on grasses located in low, wet swales. They required great effort to collect; that is, one had to sweep these grasses for hours to cap- ture only a specimen or two. I suspect the reason for this is that this species may also be living near the soil in the grass matts. Froeschner (1988c) did not list this species from North Dakota, or from any of the surrounding states or provinces. Maw et al. (2000) recorded this species from Manitoba. There is a photograph of a specimen of this species posted on the internet on BugGuide (http://bugguide.net/node/view/430768/bgimage ; accessed 31 August 2012) that was collected in Ward Co.: near Lake Darling, 11-VII-2010. 354 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Specimens Examined. MINNESOTA [New State Record]: Clay Co.: Bluestem Prairie, 9-VIII-1995, D. Rider, G. Fauske, & C. Locken (1♀ NDSU). NORTH DAKOTA [New State Record]: Foster Co.: Carrington Expt. Station, 28-IV-1988, D. D. Kopp & H. J. Meyer (1♂ NDSU). Grand Forks Co.:Oakville Prairie, Sec. 16, 47°53.403N, 97°18.855W, 21-VIII-2009, D. A. Rider (1♀ DARC), 24-VIII-2009, D. A. Rider (1♂ 1♀ DARC); Oakville Prairie, T151N, R52W, Sec. 16, 8-IX-1961 (1♂ UNDG), 26-IX-1983 (1♂ UNDG). Chlorochroa faceta (Say, 1825) - As mentioned above, there are only two species in North Dakota which have a pale, longitudinal stripe down the dorsal midline; this species can be separated from C. belfragii by the lack of black markings on the distal end of the embolium. This species is a member of the subgenus Rhytidolomia, and is nearly identical in appearance to C. lineata Thomas, a member of the nominate subgenus. Separation of these two species can be accomplished only by examining the male genitalia. Fortunately, C. lineata does not occur in North Dakota (it has been recorded only from California, Nevada, and Utah). Uhler (1872) recorded this species simply from “Dakota” without more precise information. Hussey (1922) presented the following North Dakota collection data: Ramsey Co.: shore of Devils Lake at the Narrows, July 21, 1920, T. H. Hubbell. Specimens Examined: NORTH DAKOTA: Foster Co.: McHenry, 15- IV-1967, J. Slough (1♀ MISU). Grand Forks Co.: Oakville Prairie, Sec. 9, 22- VI-2012, D. A. Rider (2♂♂ 1♀ DARC), 14-VIII-2009, D. A. Rider (2♂♂ DARC); Oakville Prairie, Sec. 9, Oakville Township, 28-VI-1961 (1♀ UNDG), 17-VII- 1961 (2♂♂ UNDG), 31-VII-1961 (2♂♂ 4♀♀ UNDG), 26-IX-1983, K. Stowe (1♀ UNDG). Slope Co.: Burning Coal Vein St. Pk., 10-VIII-1993, D. A. Rider (1♀ DARC); 11-VII-1986, K. Larsen (1♀ MISU). Ward Co.: Kenmare, 7-V-1936 (1♀ MISU); Minot, 10-V-1936, Hovdestad (1♂ MISU). Chlorochroa ligata (Say, 1831) - In most of its range, this species is relatively easy to identify; it is often a much darker, green, sometimes approach- ing black. But in northern regions, including North Dakota, they are usually the same bright green as other Chlorochroa species. Male specimens are easily identified by the male genitalia (they have a small tumescence on the posteroventral surface of the pygophore; this tumescence is lacking in other species; it is replaced in other species by a more or less obtuse, transverse, U-shaped ridge); female specimens are nearly impossible to separate from some other species, including C. persimilis, which is common in North Dakota. Lugger (1900) recorded this species from Minnesota, but at that time, workers had not realized that eastern U.S. populations were a separate species from the western species. His brief description and figure matchC. persimilis, and undoubtedly are of that species. There are no published records of this species from North Dakota. In going through material for this project, I located 4 specimens with 1981 Don Thomas determination labels for this species, but he (Buxton et al. 1983) did not include any North Dakota localities on his distribution map. He did include a locality in western South Dakota (Harris 1937, also gave a record from South Dakota). Three of the four specimens are females. One specimen, however, was a male specimen, thus verifying that C. ligata probably does occur in this state. More recently Scudder and Thomas (1987) extended the distribution of ligata into Saskatchewan and Manitoba, further lending credence to the North Dakota records. This is a western U.S. species coming into the western half of our state. Fish and Alcock (1973) studied the behavior of this species. Specimens Examined: NORTH DAKOTA [New State Record]: Oliver Co: Sec. 6, T143N, R83W, 16-VIII-1972, P. B. Kannowski (1♂ UNDG). Female specimens identified by Don Thomas in 1981: Mercer Co.: Sec. 9, T144N, R85W, July 18-20, 1972, A. C. F. Hung (1♀ UNDG). Walsh Co.: S. Branch, Park River, NE of Adams, 19-VIII-1970, J. Singleton (1♀ UNDG - this is probably C. persimilis). 2012 THE GREAT LAKES ENTOMOLOGIST 355

Chlorochroa persimilis Horváth, 1908 - Basically, there are three species of Chlorochroa in North Dakota that are bright green in color, and lack the pale stripe down the dorsal midline. Chlorochroa uhleri (see below) can be separated from the other two by having three relatively large white spots along the basal margin of the scutellum; also, the embolium is nearly as narrow apically as it is basally. Both C. ligata and this species lack the pale spots along the basal margin of the scutellum (or they are very small), and the embolium is distinctly wider apically than it is basally. To separate this species from C. ligata, please see comments under C. ligata; in short, it is nearly impossible to separate female specimens, and to separate male specimens, one must examine the pygophore. McPherson (1982) gave the distribution for this species as Québec and New England south to Florida, and west to at least the Mississippi River, and probably to Iowa and Kansas. Froeschner (1988c) listed this species from North Dakota and more eastern and southern states. Maw et al. (2000) recorded this species from Manitoba. Lugger’s (1900) record of C. ligata from Minnesota is actually a misidentification of this species. Specimens Examined: NORTH DAKOTA: Benson Co.: IX-1958 (1♂ NDSU). Billings Co.: Medora, 47°02.00N, 103°36.00W, 24-VI-2002, A. Joshi (1♀ NDSU); Roosevelt Natl. Park, Medora, 24-VI-2002, F. Antwi (1♂ NDSU); 14-VII-1965 (1♀ NDSU). Cass Co.: Fargo, 19-VI-1971, R. L. Post (1♀ NDSU). Dunn Co.: Killdeer Mts., 11-VI-1965, L. Grochowski (1♀ NDSU). Eddy Co.: 22-VIII-1958, S. Wilson (1♀ NDSU). Golden Valley Co.: T-140-104-4, D. A. Mundal, 1984, on Euphorbia esula (1♀ NDSU). Grand Forks Co.: Arvilla, 28-IX-1931 (1♀ UNDG); Emerado, 1-VIII-1949 (1♂ UNDG); Grand Forks, 10- VII-1957 (1♀ NDSU), 12-X-1964 (1♀ UNDG); Oakville Prairie, Sec. 9, T151N R52W, 15-IX-1975 (1♀ UNDG), Sec. 16, 47°53.403N, 97°18.855W, 22-VI-2012, D. A. Rider (2♀♀ DARC), 21-VIII-2009, D. A. Rider (1♂ 2♀♀ DARC), 10-IX-1958 (1♂ 1♀ UNDG); UNDG, 15-VIII-2008, B. Stargardt (1♀ UNDG); 2-IX-2008, A. Morrone (1♀ UNDG), 18-IX-2008, S. Teske (1♂ UNDG). McHenry Co.: Towner- Round Lk., VIII-1930, N. A. Weber (1♀ UNDG). McKenzie Co.: Arnegard, 5-IX-2004, E. Bruins (1♂ NDSU); 148-99-34-120, 24-VIII-1976 (1♀ NDSU). McLean Co.: Riverdale, 26-V-1977, E. U. Balsbaugh (1♀ NDSU). Mercer Co.:10-VIII-1958, Boeshans (1♀ NDSU). Mountrail Co.: Stanley, 14-IX-1998, A. Nelson (1♀ MISU). Richland Co.: 6-VIII-1958, S. Wilson (1♂ 1♀ NDSU). Slope Co.: Burning Coal Vein St. Pk., 7-VI-1973, T. L. McCabe (1♀ NDSU), 10-VIII-1993, D. A. Rider (1♀ DARC). Walsh Co.: S. Branch, Park River, NE of Adams, 19-VIII-1970, J. Singleton (1♀ UNDG). Ward Co.: Kenmare, 21-IX- 1965, M. Rutter (2♀♀ UNDG); Minot, 9-V-1986, K. Larson (1♀ MISU). Wells Co.: 11-VIII-1960, G. Mulkern (2♂♂ 1♀ NDSU). Chlorochroa uhleri Stål, 1872 - In general, this species can be recognized by the somewhat larger white spots along the base of the scutellum, and the embolium is at most slightly wider apically than it is at the base. Hussey (1922) reported this species from two North Dakota localities: Bottineau Co.: Bottineau, T. H. Hubbell, and Nelson Co.: shore vegetation behind the mud flats at Spring Lake, July 25, 1920, T. H. Hubbell. This is a western U.S. species, coming into our state from the west. Specimens Examined: NORTH DAKOTA: Billings Co.: Freeman’s Frontier near Medora, 9-VIII-1993, D. A. Rider, collected on Chenopodiaceae (1♀ DARC); Theo. Roos. Mem. Pk., 30-VIII-1961, R. L. Post (1♀ NDSU), 7-IX-1939, R. L. Post & N. M. Post (1♂ NDSU). Emmons Co.: 14-VIII-1941, J. A. Munro (6♂♂ 64♀♀ NDSU). Golden Valley Co.: Beach, IX-1921, R. L. Webster (1♀ NDSU); Sentinel Butte, 31-VIII-1961, J. Onsager & G. B. Mulkern (1♀ NDSU); VII-1958 (1♀ NDSU). Grand Forks Co.: Grand Forks, 2-X-1939 (1♀ UNDG), 3-X-1928 (1♂ UNDG); VII-1957 (1♀ NDSU). Hettinger Co.: New England, 27-XI-1940, H. S. Telford & F. P. Gutcher (2♂♂ 2♀♀ NDSU). McKenzie Co.: 14-VIII-1958, S. Wilson (1♀ NDSU). Slope Co.: Burning Coal Vein, 6-VI-1973, T. L. McCabe (1♀ NDSU), 29-VI-1972, T. L. McCabe (1♀ NDSU). Williams 356 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Co.: 21-IV-1961, J. R. Dogger (2♀♀ NDSU), 11-VII-1961, V. Goodfellow, swept from wheat (5♂♂ 11♀♀ NDSU); Williston, 12-VIII-1961, K. Brevik (1♀ NDSU). Chlorochroa viridicata (Walker, 1867) - This is a much smaller species than the other dakotan Chlorochroa species, and it is more tear-dropped shaped than other species. Neither Uhler (1878) nor Hussey (1922) reported this species from North Dakota. Froeschner (1988c) did not list this species form North Dakota either; the closest records they listed came from Montana and Nebraska. Specimens Examined: NORTH DAKOTA [New State Record]: Cass Co.: 1958 (1♂ NDSU). Slope Co.: Burning Coal Vein St. Pk., 10-VIII-1993, D. A. Rider (1♂ DARC), 12-VIII-1993, D. Cuthrell (1♂ NDSU).

Tepa Rolston and McDonald, 1984 Species of Tepa at one time were included in the genus Thyanta; in fact, Rolston (1972a), along with species now placed in Cyptocephala, called them the small Thyanta species. He provided a key for their identification; then later (Rolston and McDonald 1984) described a new genus, Tepa, to hold these species. Rider (1986b) made a couple corrections, and provided a new key for their identification. There are currently sixTepa species known, two of which are known to occur in North Dakota, and a third species has been recorded from South Dakota. The following key will separate our two species, as well as T. vanduzeei Rider in case it might eventually be found in North Dakota.

Key to species of Tepa that may be found in North Dakota 1 Ostiolar ruga elongate, distance from mesial margin of ostiole to apex of ruga much greater than distance from apex of ruga to lateral margin of supporting sclerite [not yet found in North Dakota]...... vanduzeei Rider – Ostiolar ruga shorter, distance from mesial margin of ostiole to apex of ruga equal or less than distance from apex of ruga to lateral margin of supporting sclerite...... 2 2(1) Anterolateral pronotal margins straight to slightly convex; ostiolar ruga nearly evanescent apically in profile...... brevis (Van Duzee) – Anterolateral pronotal margins distinctly concave; ostiolar ruga truncate apically in profile...... rugulosa (Say) Tepa brevis (Van Duzee, 1904) - The two species of Tepa known from North Dakota are relatively easy to separate. This species has the anterolateral pronotal margins substraight; in T. rugulosa, they are distinctly concave. This is primarily a western or southwestern species, coming in the western third of North Dakota. There are no published records of this species from North Dakota. Neither Froeschner (1988c) nor Maw et al. (2000) listed this species from North Dakota or any surrounding states or provinces. The closest known record is from Colorado (Rider 1986b). Some of the specimens collected listed below were collected on a low bushy plant in the Chenopodiaceae. Specimens Examined: NORTH DAKOTA [New State Record]: Bill- ings Co.: Freeman’s Frontier near Medora, 9-VIII-1993, D. A. Rider, on Che- nopodiaceae (3♂♂ 2♀♀ DARC); Freeman’s Frontier, T140N, R101W, Sec. 25, 9-VIII-1993, G. Fauske (1♀ NDSU). Tepa rugulosa (Say, 1831) - In this species, the anterolateral pronotal margins are distinctly concave, whereas members of T. brevis have the margins substraight. The lectotype and at least one paralectotype (see Rider 1986b) of Thyanta punctiventris Van Duzee were collected by Prof. Wickham from 8-9 June, and are from Williston (Williams Co.), North Dakota. Thyanta punctiventris is a synonym of this species. This is a western or southwestern U.S. species, coming 2012 THE GREAT LAKES ENTOMOLOGIST 357 only into the western third of North Dakota. Some specimens listed below were collected with the previous species on a small bush in the family Chenopodiaceae. Specimens examined: NORTH DAKOTA: Billings Co.: Freeman’s Frontier near Medora, 9-VIII-1993, D. A. Rider, on Chenopodiaceae (1♂ 5♀♀ DARC). Slope Co.: Burning Coal Vein St. Pk., 10-VIII-1993, D. A. Rider, at light (1♀ DARC). Williams Co.: Williston, 23-VIII-1987, M. J. Weiss (1♂ NDSU).

Thyanta Stål, 1862 Thyanta is a relatively large genus, and most species occur in the Neo- tropical regions; there are seven species that occur in North America, but three of these are tropical just reaching into the southern areas of the U.S.; a fourth species is endemic to southern Florida, and a fifth species is western reaching only as far east as Utah. The other two species are widely distributed in the eastern U.S., but one of these, T. calceata (Say), does not reach North Dakota. Only one species occurs in North Dakota. The genus was recently revised in two parts; the first part covered the South American species (Rider and Chapin 1991), while the second part covered those species occurring north of South America (Rider and Chapin 1992). Members of this genus tend to be small to medium in size, usually pale to bright green (there is some yellow to brown autumnal forms), sometimes with reddish or blackish markings; the ostiolar rugae are elongate, tapering to an acuminate apex. Thyanta custator accerra McAtee, 1919 - The nominate subspecies occurs along the Gulf and Atlantic coasts, and is distinguished by black markings along the anterolateral pronotal margins, and a small black spot located in the inner angle of each pronotal cicatrice (both are lacking in this subspecies). The distribution given by McPherson (1982) was New York south to Georgia, and west to Idaho, northeastern Utah, northeastern New Mexico, and Texas. Uhler (1876) listed this species simply from ‘Dakota’; this may serve as the basis for Froeschner (1988c) listing it from North Dakota. Hussey (1922) did not record it from North Dakota. The only published records are those of Rider and Chapin (1992), who listed it from three North Dakota counties (Cass Co., Hettinger Co., and Stark Co.) without any specific collecting data. Specimens Examined: NORTH DAKOTA: Cass Co.: Fargo, 915 11th Ave. N., 10-X-2008, J. Sanden (1♀ NDSU); Fargo, NDSU campus, 6-IX-2001, D. A. Rider (1♂ DARC), 9-IX-2003, B. Dokter (1♂ NDSU); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 1-VIII-2012, D. A. Rider (1♀ DARC); 7-VI-1940 (1♀ NDSU). Hettinger Co.: 9-VIII-1955, V. Goodfellow (1♀ NDSU). McIntosh Co.: Wishek, 12-IX-2005, B. Aipperspach (1♂ NDSU). Ransom Co.: SNG, A Annex, 19-V-2005, G. M. Fauske, at light (1♂ NDSU). Richland Co.: Ekre Grassland Preserve, 48°32′15″N, 97°12′00″W, 8-VII-2002, A. Joshi (1♀ NDSU). Slope Co.: Amidon, 23-VI-2002, J. Knott (1♂ NDSU). Ward Co.: Minot, 20- VII-2002, G. Hanley (1♀ MISU); NCREC, Minot, 13-VII-1999, A. Culbertson, blacklight trap (1♀ NDSU).

Tribe Pentatomini Leach, 1815 This is a difficult tribe to define. Basically, after most other genera have been placed in other tribes, those remaining genera which have a spine or tubercle at the base of the abdomen have been placed in the Pentatomini. The basal abdominal spine or tubercle is not confined to the Pentatomini however. This character also occurs in the Menidini, some Nezarini, some Procleticini, and others. The type of the tribe, Pentatoma, is an Old World genus that is much larger, and very differently colored than Banasa. So, its placement in the Pentatomini is tentative at best; it may be better suited in the Menidini, near the genus Rio. 358 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Banasa Stål, 1860 Banasa is a very diverse New World genus with most of its species occurring in the Neotropics. Twelve species are recorded from North America, but only two occur in our area. Banasa was revised in three parts: the first part covered North America (Thomas and Yonke 1981), the species in Mexico, Central America, and the Antilles were treated in the second part (Thomas and Yonke 1988), and the South American species were covered in the third part (Thomas and Yonke 1990).

Key to species of Banasa in North Dakota 1 Dorsal surface bright green with distinct white mottled spots; pronotal disk monocolorous...... euchlora Stål – Dorsal surface colored differently; pronotal disk usually bicolored, posterior half much darker than anterior half...... dimidiata (Say) Banasa dimidiata (Say, 1832) - This species can be recognized by the distinctly bicolored pronotum (see key above). Say originally spelled the species name dimiata, but LeConte emended the name to dimidiata in his reproduction of Say’s works. Froeschner (1988c) considered this an unjustified emendation, but Hoffman (2005) argued for the continued use of the emended form dimidiata. McPherson (1982) gave the distribution as Québec and New England west to the Pacific coast, and south to Florida, Oklahoma, Texas, and California. Although he gave no specific localities, Van Duzee (1904) reported that he had examined specimens of this species from North Dakota. This may serve as the basis for Froeschner (1988c) also listing it from North Dakota. Decoursey (1963) described the life history of B. dimidiata. This species has been collected on several bushes or trees including juniper and ash. Specimens Examined. NORTH DAKOTA: Billings Co.: Th. Roos. M. Prk., 14-VII-1965, W. Kotchman (1♂ NDSU). Cass Co.: Absaraka Arboretum, 20-IX-2002, S. Koudela (1♂ NDSU); NDSU Campus, 25-VI-2012, D. A. Rider (3♀♀ DARC), 4-IX-2004, C. Bye (1♂ NDSU), 4-IX-2008, D. A. Rider (2♂♂ 2♀♀), 6-IX-2000, S. Kveno (1♂ NDSU), 6-IX-2001, D. A. Rider (1♂ 3♀♀ DARC); W NDSU Campus, 21-V-2012, D. A. Rider (1♂ DARC); n. Fargo, 1-VIII-2012, C. Erhard (1♀ NDSU); ne Fargo, 46.759N, 96.765W, 31-VII-2012, D.Rider (1♂ 1♀ DARC); Wild Rice R. at Univ. Dr., 46.759N, 96.797W, 27-VII-2012, D. A. Rider (1♂ DARC), 13-VIII-2012, D. A. Rider (1♂ 1♀ DARC); V-1963, G. Nitz (1♀ UNDG); IX-1957 (1♂ NDSU). Grand Forks Co.: State Park near Arvilla, 1-X- 1969 (1♂ UNDG); Grand Forks, 4-X-1961 (1♀ UNDG), 27-X-1993 (1♂ UNDG); 20mi. S. Grand Forks, 19-IX-1970 (1♀ UNDG); Oakville Prairie, Secs. 9&26, T151N, R52W, 23-IX-1968, D. Klein (1♂ UNDG); Turtle R., 4.8mi. W., 2mi. N. Emerado, T152N, R53W, Sec. 20 & 29, 22-IX-1979 (1♂ UNDG), 24-IX-1976 (1♂ UNDG), 28-X-1980 (1♀ UNDG). Morton Co.: Mandan, 22-VII-1974, P. K. & B. A. Lago (1♂ NDSU), 22-IX-1922, R. L. Webster (1♂ NDSU). Nelson Co.: McVille, 1-IX-2004, J. Stein (1♀ NDSU). Ramsey Co.: Grahms Island State Park, 28-V-2007, T. Retterrath, Lindgren trap (1♂ NDSU). Ransom Co.: Lisbon, 14-VII-1983, D. Siebert (2♂♂ 2♀♀ NDSU); 3mi N of Mirror Pool, T135N R53W, Sec 2, 13-VI-1992, D. Cuthrell (2♀♀ NDSU); 4mi. NE Sheldon, 19-VI-1984, M. McLeod (1♂ 1♀ NDSU); Walcott Dunes, 13-IX-1994, M. Ellefson (1♀ NDSU). Richland Co.: Ekre Grassland Presv., 25-IX-1994, T. Nowatzki (1♂ NDSU); Ekre Grassland Preserve, T136N, R51W Sec. 32, 6-IX-1994, K. Urlacher (1♂ NDSU); Mirror Pool, T135N R52W Sec 8, 3-VI-1994, D. A. Rider (4♀♀ DARC), 10-VIII-1994, J. L. Jyoti (1♂ NDSU), 28-VIII-1994, J. L. Jyoti (1♀ NDSU), 29-VIII-1996, D. Rider (2♀♀ DARC), 6-IX-1994, T. Nowatzki (1♂ NDSU); 1.5mi NE Mirror Pool, 15-VI-1994, D. A. Rider (3♀♀ DARC); SNG, Oak Savannah, T135N R52W Sec. 10, 26-VI-1996, L. DeCock, Hg vapor light 2012 THE GREAT LAKES ENTOMOLOGIST 359 (1♀ NDSU), 26-VI-1996, L. Pederson (1♂ NDSU). Walsh Co.: 1mi. E of Pisek, 24-VII-1976 (1♂ UNDG); S. Branch, Park River, NE of Adams, 19-VIII-1970, J. Singleton (1♀ UNDG). Banasa euchlora Stål, 1872 - This is one of the easiest Banasa species to identify; the bright green coloration with relatively large pale spots on the dorsum is diagnostic. Froeschner (1988c) did not list this species from North Dakota, or from any of the surrounding states or provinces (closest records included Iowa and Illinois). This species is very common in the southern and southwestern United States, and becomes rather sparse and sporadic farther north. All known North Dakota records are from the southern half of the state. There is also a ♀ specimen in the NDSU Collection from South Dakota, Fall River Co.: Bttl. Mt. Game Prod. Area, 7-VIII-1984, G. Fauske, which appears to represent a new state record. Specimens Examined: NORTH DAKOTA [New State Record]: Burleigh Co.: Bismarck, 6-XII-2010, T. Quanrud, inside home (1♀ NDSU). Cass Co.: Fargo, NDSU, 2-X-1992, S. Cong, at light (1♀ NDSU), 9-X-2002, J. Dykshoorn (1♂ DARC). Richland Co.: VII-1957 (1♂ NDSU).

Tribe Sciocorini Amyot and Serville, 1843 This is primarily an Old World tribe with only a few genera, but many species. There are only two sciocorine genera in the New World. Antillosciocoris Thomas contains a single species known only from the Dominican Republic in the West Indies. The South American genus Trincavellius Distant has been placed in the Sciocorini, but it probably belongs in another tribe. Sciocoris is the only sciocorine genus known from the North America. Members are fairly small, and usually have the lateral margins of the head, pronotum, and/or corium expanded, somewhat explanate. A single genus with a single species occurs in our region.

Sciocoris Fallén, 1829 This is primarily an Old World genus with only three New World species known; in fact, the species in our area is Holarctic, occurring also in northern parts of the Old World. The other two New World Sciocoris species are S. crassus Ruckes (occurs in Mexico) and S. longifrons Ruckes (occurs in Texas) . Members of the Sciocorini tend to be smallish, orbicular, and usually have the lateral margins of the head, pronotum, and coria somewhat explanate. Sciocoris microphthalmus Flor, 1860 - This is the only species of this genus that occurs in North Dakota. The small, ovate shape coupled with the explanate pronotal margins will separate this species from all other North Da- kota’s pentatomids. McPherson (1982) gave the distribution as Québec west to Alberta, and south to Massachusetts, Iowa, North Dakota, and possibly Okla- homa. Hussey (1921, 1922) recorded this species from Slope Co.: rocky side of Black Butte, near Amidon, August 21, 1920, T. H. Hubbell. Specimen examined: NORTH DAKOTA: Grand Forks Co.: Oakville Prairie, T151N, R52W, Sec. 16, 10-IX-1980, TLS (1♀ UNDG).

Tribe Strachiini Mulsant and Rey, 1866 This tribe contains more genera and species in the Old World than in the New World, but there are several western hemisphere genera. Its included genera are characterized by the ostiole being very small, obsolete, and located more mesially between the mid and hind coxae; the ostiolar rugae and associated evaporative areas are also obsolete. Many of these species are brightly 360 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4 colored with reds, oranges, and/or blues. Until recently, only one genus occurred in North America, Murgantia. In the past five years, specimens of Bagrada hilaris have been discovered in California and Arizona (Palumbo and Natwick, 2010). These are much smaller, and much less brightly colored than Murgantia. In general, species of this tribe seem to feed on members of the Brassicaceae.

Murgantia Stål, 1862 This New World genus, at present contains a number of species mostly of neotropical distribution. A single species, M. histrionica (Hahn), is common throughout much of the United States. There is, however, some evidence that M. histrionica may not be congeneric with the other species placed in this genus. Again, this will require further study. Murgantia histrionica (Hahn, 1834) - This is a very easily recognized species with its bright orange coloration on a shiny, black background. Parshley (1922) listed this species from South Dakota, and Froeschner (1988c) listed it from Minnesota; I have not located the source of that record. There is a single specimen in the UNDG collection from North Dakota; it is from Grand Forks Co., or about two-thirds up the Red River Valley. Its presence in North Dakota needs verification (the specimen listed below appears to have been collected by a student as the label was untrimmed, there is no specific data, and there was also a “Pentatomidae” label). Additionally, Lugger (1900) reported that M. histrionica had not been found in Minnesota. The biology of this species has been studied (Canerday 1965), and more recently, McPherson and Ahmad (2008) described the genitalia. Specimen Examined: NORTH DAKOTA [New State Record]: Grand Forks Co.: 8-IX-2008, S. Teske (1♂ UNDG).

PENTATOMIDAE: PODOPINAE Amyot and Serville, 1843 Not only is this a relatively small subfamily in numbers (not many genera or species), but also in size as most of its members are relatively small. This subfamily is much more diverse in the Old World. Most species are rather drab gray or brown to black in color, but a few species in the Old World are conspicuous red with black markings (Graphosoma). They are commonly called turtle bugs, perhaps because of their turtle-like shape, or because of the fact that some species are commonly found on grasses near wet marshy areas. In fact, several species are known to be considerable pests of rice in both the Old World (Scotinophara) and New World (Amaurochrous). Several genera occur throughout North America, but only one, Amaurochrous, has species known from North Dakota.

Amaurochrous Stål, 1872 This genus contains six North American species, but two of these occur only in the very eastern parts of the U.S.; one is only known from California; and one is widespread in the eastern states, but does not reach North Dakota. This leaves two species which are known to occur here. A review of this genus was provided within the review of the entire subfamily for North America (Barber and Sailer 1953). Key to species of Amaurochrous in North Dakota 1 Anterolateral pronotal margins distinctly sinuous, distinctly concave along posterior two-thirds; head relatively narrowly rounded, width at apex at most slightly wider than width just in front of eyes; juga slightly longer than tylus, but usually not contiguous...... cinctipes (Say) 2012 THE GREAT LAKES ENTOMOLOGIST 361 – Anterolateral pronotal margins slightly sinuous, nearly straight, no where distinctly concave; head broadly rounded apically, distinctly wider near apex than width just in front of eyes; juga distinctly longer than tylus and usually contiguous anteriorly...... brevitylus Barber and Sailer Amaurochrous brevitylus Barber and Sailer, 1953 - This species can be separated from A. cinctipes by its nearly straight anterolateral pronotal margins. There are no published records of this species from North Dakota although Froeschner (1988c) listed it from Minnesota. We have collected many specimens of this species from Bluestem Prairie in western Minnesota (Clay Co.). Specimen Examined: NORTH DAKOTA [New State Record]: Cass Co.: X-1957 (1♂ NDSU). Amaurochrous cinctipes (Say, 1828) - This species can be separated from A. brevitylus by the distinctly sinuous anterolateral pronotal margins. There has been no report of this species from North Dakota; Froeschner (1988c) listed it from Minnesota. The ecology and biology of this species has been studied (Parshley 1923, Davis 1925). McPherson and Paskewitz (1984b) reared this species in the laboratory, and described the immature stages. Specimens Examined: NORTH DAKOTA [New State Record]: Rich- land Co.: Ekre Grassland Preserve, 3-IX-2009, D. A. Rider (1♂ DARC); 19-VI- 1966 (1♀ NDSU).

SCUTELLERIDAE Leach, 1815 Members of the Scutelleridae lack the spines on the tibiae characteristic of the Cydnidae and Thyreocoridae; they have three tarsal segments, similar to the Pentatomidae, but the scutellum is usually greatly enlarged, covering nearly the entire abdominal dorsum (if the scutellum is somewhat spatulate, then the exposed portion of the corium narrows apically). The family can be divided into several subfamilies, three of which have species occurring in North Dakota. The North American scutellerid species have been treated in an unpublished dissertation (Lattin, 1964).

Key to the Subfamilies of Scutelleridae in North Dakota 1 Scutellum spatulate, corium exposed for nearly its entire length, becoming somewhat acuminate apically...... Eurygastrinae – Scutellum more broadly rounded, covering most of abdomen, corium only exposed basally, apex angulate...... 2 2(1) Abdominal sternites IV and V with a series of very small ridges forming stridulatory areas, these ridges can be difficult to see without a proper microscope and light system...... Pachycorinae – Abdominal sternites IV and V lacking stridulatory areas...... Odontotarsinae

SCUTELLERIDAE: EURYGASTRINAE Amyot and Serville, 1843 Members of this tribe can be recognized by the more spatulate scutellum that does not cover the entire abdomen; that is, the corium is exposed along the lateral margins for its entire length. The Eurygastrinae contains only a few genera, most of which are restricted to the Old World in distribution. The only genus known from the New World, Eurygaster, has most of its species distributed in the Eastern Hemisphere. 362 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Eurygaster Laporte, 1833 There are several species of Eurygaster described from North America, but only two, E. alternata (Say) and E. amerinda Bliven, may occur in our area; only E. alternata has been recorded from North Dakota. Eurygaster amerinda has been recorded from California and Illinois (Lattin, 1964) and Manitoba and Saskatchewan (Maw et al., 2000). Eurygaster amerinda has the anterolateral pronotal margins distinctly convex, and the apex of the head broadly rounded. In E. alternata, the anterolateral pronotal margins are nearly straight, and the apex of the head is more narrowly rounded. In the material listed below, there is an occasional specimen that has the anterolateral pronotal margins slightly convex and perhaps the head is slightly broader, but I do not see any other significant differences, especially in the male or female genitalia. I have chosen to consider all specimens I have examined from North Dakota to be E. alternata until a more thorough study can be done. The North American species of Eurygaster were reviewed by Vodjani (1961). Eurygaster alternata (Say, 1828) - McPherson (1982) gave the range for this species as across the entire northern part of the continental United States and southern Canada, but also ranges southward to Virginia and New Mexico. Uhler (1878) recorded this species from three different North Dakota localities: Bottineau Co.: prairies near Mouse River, August 29, Pembina Co.: near Pembina, and Rollette Co.: Turtle Mountains. Hussey (1922) added three more records from two additional counties: Ramsey Co.: Devils Lake, July 25-28, 1920, T. H. Hubbell, Slope Co.: Chalky Buttes south of Amidon, August 22, 1920, T. H. Hubbell and Amidon, edge of wheat field, August 24, 1920, T. H. Hubbell. Specimens Examined: NORTH DAKOTA: Billings Co.: Easy Hill, T138N, R100W, D. Aarhus (1♂ 1♀ NDSU); Freeman’s Frontier nr. Medora, 9-VIII-1993, D. A. Rider (1♂ DARC); Medora, 47 02 00N, 103 36 00W, 24-VI- 2002, A. Joshi (1♂ 2♀♀ NDSU); TR Elkhorn Rch, 25-VI-1965, W. Kotchman (1♀ NDSU). Cass Co.: 10-X-2003, R. Wagner (1♀ NDSU). Dunn Co.: Killdeer Mts., 11-VI-1965, L. Grochowski (1♀ NDSU), W. Kotchman (1♂ 1♀ NDSU), 28-VI-1965, L. Grochowski (1♀ NDSU), 3-VIII-1965, W. Kotchman (1♀ NDSU); 21-VI-1977, L. A. Schutz, J. W. Smith (♀ NDSU), 3-VII-1966 (1♀ NDSU), 13- VII-1976 (1♂ NDSU). Grand Forks Co.: Oakville Prairie, Sec. 16, 47°53.403N, 97°18.855W, 22-VI-2012, D. A. Rider (1♂ 2♀♀ DARC), 24-VIII-2009, D. A. Rider (1♂ 1♀ DARC). McKenzie Co.: Th. Roos. M. Park, 22-VI-1965, W. Kotchman (1♂ NDSU), 24-VI-1964, R. L. Post (1♀ NDSU), 20-VII-1965, W. Kotchman (1♂ NDSU), L. Grochowski (♀ NDSU); 13-VII-1976, Lago & Kurtz (1♀ NDSU). McLean Co.: 18-VII-1958, S. Wilson (1♀ NDSU). Ransom Co.: 2mi. SW McLeod, T134N, R53W, sec. 35, 9-VI-1992, D. A. Rider (1♀ DARC); 4mi. NW of McLeod, T134N, R53W, Sec. 6, 18-VI-1992, D. A. Rider (1♀ DARC). Richland Co.: 0.5mi. SE Mirror Pool, 6-VI-1995, D. A. Rider (1♀ DARC); SNG, Oak Sa- vannah Area, 18-VI-2000, D. A. Rider (1♂ 1♀ DARC), 17-VII-1993, D. Cuthrell (1♀ NDSU); SNG, Horsetrail Head, 46°31.409N, 97°12.194W, 28-V-2010, D. A. Rider (3♂♂ 6♀♀ DARC), 27-VI-2012, D. A. Rider (1♂ 1♀ DARC), 20-VII-2012, D. A. Rider (1♂ DARC), 25-VIII-2009, D. A. Rider (4♂♂ 2♀♀ DARC), 4-IX-2009, D. A. Rider (1♀ DARC). Slope Co.: Burning Coal Vein St. Pk., 26-VI-1973, T. L. McCabe (1♀ NDSU), 10-VIII-1993, D. A. Rider (2♀♀ DARC); 9-VI-1977 (1♂ 1♀ NDSU). Stutsman Co.: Chase Lake, 4-VI-1973, T. L. McCabe (1♂ 1♀ NDSU). Ward Co.: Minot, 6-VII-1986, K. Larsen (1♀ MISU).

SCUTELLERIDAE: ODONTOTARSINAE Mulsant and Rey, 1865 Scutellerids with a more spatulate scutellum are placed in the Eurygastrinae (see above); the remaining genera with a more extensive scutellum (corium only exposed basally) can be split into two subfamilies. Those which have a series of stridulatory ridges on the abdominal venter belong in the Pachycorinae (see below), 2012 THE GREAT LAKES ENTOMOLOGIST 363 and those genera which lack the stridulatory ridges belong in this subfamily. These genera were treated in the Eurygastrinae by McPherson (1982). Again, many of the included genera occur mainly in the Old World, but several are known from North America. Members in this subfamily tend to not be collected as commonly as members of the other two subfamilies. There are three Odontotarsine genera which occur, or may occur, in North Dakota; they can be separated by the following key.

Key to the Odontotarsinae Genera that may occur in North Dakota (modified from Lattin 1964) 1 Anterolateral pronotal margins strongly sinuate...... 2 – Anterolateral pronotal margins substraight, at most only slightly sinuous...... Vanduzeeina 2(1): Anterior margin of pronotum produced over base of head; tylus and juga strongly developed, protuberant [not yet recorded from North Dakota]...... Euptychodera – Anterior margin of pronotum not produced over base of head; tylus slightly elevated above juga, but not strongly developed or protuberant...... Phimodera

Euptychodera Bergroth, 1908 This New World genus contains a single species, E. corrugata (Van Duzee). Its distribution is south and west of North Dakota, but Harris (1943) recorded it from South Dakota. Because this species is not common, I am providing the locality data from the single specimen housed in NDSU from South Dakota: Yankton Co.: Yankton, 17-V-1967, V. M. Kirk, beach (1♂ NDSU). It is possible that this species may eventually be found in North Dakota.

Phimodera Germar, 1839 This genus can be recognized by the lack of stridulatory ridges on the abdominal venter, the distinctly sinuous anterolateral pronotal margins, and the anterior pronotal margin not produced over the base of the head. This genus presents an interesting taxonomic problem; three species (with several subspecies) have been described from North America, but Lattin (1964) in his unpublished thesis placed P. torpida Walker and P. torrida Reuter as synonyms of P. binotata (Say). It does not appear that this synonymy has been published, and so all three names are still in use (all three were cataloged as valid species in Froeschner 1988d). This does not affect this manuscript greatly as only the oldest named species has been recorded from North Dakota. But P. torpida was listed from Saskatchewan and Michigan by Froeschner (1988d); if it is a valid species, then it could possibly be found in North Dakota, if it is a synonym, then it fits with the known distribution ofP. binotata. Phimodera binotata (Say, 1824) - The distribution of this species was given by McPherson (1982) as Manitoba and Michigan west and southwest to British Columbia, Washington, Oregon, California, Arizona, and New Mexico. There are no published records of this species from North Dakota, although Froeschner (1988d) listed it from Manitoba and South Dakota, and Maw et al. (2000) recorded it from Manitoba and Saskatchewan. Specimens Examined: NORTH DAKOTA [New State Record]: Ransom Co.: SNG, T135N, R53W, Sec. 29, 13-VIII-2000, J. Maxwell, collected on sand hills nr. Hudsonia (1♀ NDSU). Richland Co.: Walcott Dune, 20-VIII-1979, J. W. Smith (2♀♀ NDSU). Ward Co.: Minot, 7-V-1936, C. Bigler (1♀ MISU), 10- VIII-1987, M. J. Weiss (3♂♂ 2♀♀ NDSU). Williams Co.: Williston, 19-V-1987, 364 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

M. J. Weiss (2♂♂ NDSU), 14-VI-1987, M. J. Weiss (1♂ 3♀♀ NDSU), 12-VII-1987, M. J. Weiss (1♀ NDSU), 26-VII-1987, M. J. Weiss (2♀♀ NDSU).

Vanduzeeina Schouteden, 1904 This genus contains five species, only one of which occurs in our region. Three are known only from California, and the fourth is western in distribution, reaching only as far east as Colorado. Usinger (1930) provided a key to the species of Vanduzeeina. Vanduzeeina borealis Van Duzee, 1925 - It is possible to confuse this species with Homaemus aeneifrons if you do not closely examine the specimens. In general, they both have a similar color pattern, and they are about the same size. This species, however, tends to have a much more robust abdomen, it lacks the stridulatory ridges on the abdomen, and it tends to be darker brown all over. McPherson (1982) gave the distribution for this species as Illinois, South Dakota, Manitoba, Alberta, British Columbia, and the Yukon Territory. Froeschner (1988d) listed this species from Manitoba and South Dakota, and Maw et al. (2000) also recorded this species from Manitoba. Apparently, this species has not previously been reported from either Minnesota or North Dakota. It is interesting to note that all the specimens from Minnesota were collected in pitfall traps; there is one more specimen in the NDSU collection from Manitoba that was also collected in a pitfall trap. Specimens examined: MINNESOTA [New State Record]: Norman Co.: Agassiz Dunes SNA, 47°29′N 96°18′W, 18-VI-1996, C. Locken, D. Rider, pitfall (2♀♀ NDSU), 1-VII-1997, A. Abbott, P. Tinerella, pitfall (1♀ NDSU), 9-VII-1999, P. Tinerella, G. Fauske, pitfall (2♀♀ NDSU), 17-VII-1996, K. Ur- lacher, C. Locken, pitfall (1♀ NDSU), 30-VII-1996, D. Rider, C. Locken, pitfall (1♀ NDSU). Polk Co.: Agassiz Dunes SNA, 47°31′N, 96°18′W, 11-VI-1998, P. Tinerella, pitfall (1♀ NDSU), 29-VI-1999, P. Tinerella, G. Fauske, pitfall (1♀ NDSU), 1-VII-1997, P. Tinerella, A. Abbott, pitfall (1♂ NDSU), 9-VII-1999, P. Tinerella, G. Fauske, pitfall (1♀ NDSU), 17-VII-1996, D. Rider, C. Locken, pitfall (1♀ NDSU), 21-VII-1997, P. Tinerella, A. Abbott, G. Fauske, pitfall (2♀♀ NDSU), 27-VII-1999, C. Davis, C. Jordan, pitfall (2♀♀ NDSU), 12-VIII-1996, K. Urlacher, D. Rider, pitfall (1♀ NDSU), 21-VIII-1997, P. Tinerella, A. Abbott, pitfall (1♀ NDSU). NORTH DAKOTA [New State Record]: Slope Co.: Burning Coal Vein St. Pk., 23-VI-1965, L. Grochowski (1♀ NDSU).

SCUTELLERIDAE: PACHYCORINAE Amyot and Serville, 1843 In our region, this subfamily contains those scutellerid genera which have a relatively extensive scutellum, covering most of the abdomen, and also has a series of stridulatory ridges on the abdominal venter. This is a fairly large subfamily with quite a few included genera. Only two genera, however, have members recorded from North Dakota; there is a chance that a species from a third genus (Tetyra) may eventually be found in North Dakota.

Key to Pachycorinae Genera that May Occur in North Dakota (modified from McPherson 1982) 1 Pronotum with distinct, transverse, submedian impression; head strongly declivent; color black...... Acantholomidea – Pronotum without distinct, transverse impression; head gradually declivent; color usually not black...... 2 2(1) Ostiole extended as distinct canal toward lateral margin of supporting plate...... Homaemus 2012 THE GREAT LAKES ENTOMOLOGIST 365 – Ostiole not extended as canal toward lateral margin of supporting plate...... Tetyra

Acantholomidea Sailer, 1945 Members of this genus are relatively distinctive with the stridulatory ridges on the abdominal venter, and the transverse pronotal impression. There are two North American species in this genus, one of which has been found in North Dakota, and is treated below. The known distribution of the other, A. porosa (Germar), seems to be more western or southern. In A. porosa, the anterolateral pronotal margins lack the teeth or denticles which are present in A. denticulata. Acantholomidea denticulata (Stål, 1870) - This species is easy to recognize by the transverse pronotal impression coupled with the denticles along the anterolateral pronotal margins. McPherson (1982) gave the distribution as Ontario, New York south to Virginia, and west to Minnesota and Kansas. There are no published records of this species from North Dakota; Froeschner (1988d) listed it from Minnesota. The biology of this species has been studied (Harris and Andre 1934). Specimen Examined: NORTH DAKOTA [New State Record]: Richland Co.: Mirror Pool, T135N, R52W, Sec. 8, NE 1/4, 23-VI-1966, Gordon & Aarhus (1♀ NDSU).

Homaemus Dallas, 1851 In our region, members of this genus can be recognized by the presence of the stridulatory ridges on the abdominal venter, and an ostiolar canal extending laterally and then curving rather abruptly anteriorad at its apex. There are five species (and several subspecies) that occur in North America. Again, some confusion exists stemming from the unpublished Lattin (1964) thesis. Lattin (1964) recognized five species, but he considered consors to be a subspecies of aeneifrons, and extensus Walley as a synonym of consors. Froeschner (1988d) treated consors as a synonym of bijugis and extensus as a subspecies of aeneifrons. This is not a major problem for this paper, however, as only two species are known from North Dakota: the nominate subspecies of H. aeneifrons and H. bijugis. The majority of specimens in our region can be readily separated by the presence (bijugis) or absence (aeneifrons) of pale submarginal stripes on the head. There is, however, a small percentage of H. bijugis with the pale stripes nearly lacking, and conversely some specimens of H. aeneifrons that have a trace or more of the stripes present. Additionally, I have found that the female abdominal sternite character (see key below) is not always reliable. There is a color character for female specimens that seems to work well. Male specimens tend to have the scutellum solid brown in color, with a few dark punctures and a few pale punctures in no apparent pattern. Female specimens tend to have a thin, pale, medial longitudinal line that tends to widen distally, forming a pale band. In H. bijugis, the thin line widens very gradually to form a relatively narrow pale band; in H. aeneifrons, the thin line widens rather abruptly, and forms a slightly wider pale band. In general, I have also found that specimens of H. aeneifrons tend to be slightly larger and a little broader, but this is not always evident unless one has a series of both species.

Key to the species of North Dakota Homaemus 1 Dorsum of head nearly uniformly black, usually lacking yellow submarginal stripe on each jugum, occasionally yellow stripe partially present; overall size and shape slightly larger, broader, more rounded; female 366 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

with last abdominal sternite flattened or concavely impressed postero- medially, the posterior margin of which forms a transverse carina just anterior to genital plates; female with thin, pale, mediolongitudinal line on scutellum abruptly widening forming a relatively broad pale band near apex...... aeneifrons (Say) – Dorsum of head black with distinct yellow submarginal stripe on each jugum, occasionally yellow stripe invaded with small amount of black; overall size and shape somewhat smaller, narrower, more elongate; female with last abdominal sternite not concavely impressed posteromedi- ally; female with thin, pale, mediolongitudinal stripe gradually widening to form a relatively narrow pale band apically...... bijugis Uhler Homaemus aeneifrons (Say, 1824) - At one time or another, this species has been split into three subspecies. Lattin (1964) considered H. a. extensis Walley to be a synonym of H. a. consors, so at present only two subspecies are recognized. The nominate subspecies occurs in our area; H. a. consors Uhler is more western in distribution. In the past, some workers have considered consors to be a valid species. McPherson (1982) gave the distribution as northeastern Nova Scotia south to Tennessee and western North Carolina, and west to Manitoba and Kansas. Uhler (1878) provided two records of this species from North Dakota: Bottineau Co.: vicinity of Mouse River, August 29, 1873, T. H. Hubbell, and Pembina Co.: near Pembina. Hussey (1922) added an additional three North Dakota records: Cass Co.: Fargo, T. H. Hubbell; Ramsey Co.: Devils Lake, July 25, 1920, T. H. Hubbell; and Rollette Co.: Turtle Moutains, T. H. Hubbell. This is species is quite common in North Dakota. Specimens Examined: NORTH DAKOTA: Barnes Co.: Valley City, 26-VIII-1917, P. W. Fattig (1♂ NDSU); T139N, R58W, Sec. 14, 18-IX-2004, P. S. Burange (1♀ NDSU). Billings Co.: TR Elkhorn Rch, 25-VI-1965, L. Gro- chowski (2♂♂ NDSU), 14-VII-1965, W. Kotchman (1♂ NDSU). Bottineau Co.: 28-VIII-1961, R. L. Post (1♂ NDSU). Cass Co.: Fargo, VIII-1971, M. Knutson (1♀ NDSU); Magnolia WMA, T140N, R54W, Sec. 25, 10-IX-1994, K. Urlacher (1♀ NDSU); Wild Rice R. at Red R., 46.758N, 96.787W, 13-VIII-2012, D. A. Rider (1♂ DARC); 18-VII-1960, D. Noetzel (1♀ NDSU), 18-VIII-1958, S. Wilson (1♀ NDSU), 27-IX-1958 (1♀ NDSU). Dunn Co.: 13-VII-1976, Lago & Kurtz (1♂ NDSU), 28-VII-1976, Lago & Kurtz (1♂ 1♀ NDSU). Grand Forks Co.: Inkster Spring, 13-VII-1964, R. J. Sauer (1♀ NDSU); Oakville Prairie, Sec. 16, 47°53.403N, 97°18.855W, 22-VI-2012, D. A. Rider (1♂ DARC), 21-VIII-2009, D. A. Rider (2♂♂ 1♀). Hettinger Co.: Indian Creek Dam, T133N, R95W, Sec. 17, 8-X-1994, K. Urlacher (1♀ NDSU). McKenzie Co.: Th. Roos. M. Prk., 21- VII-1965, L. Grochowski (1♂ NDSU); 27-VII-1976, Lago & Kurtz (1♂ NDSU); 10-VIII-1976 (1♀ NDSU), 24-VIII-1976 (1♀ NDSU). McLean Co.: 10-VI-1957, P. Bergen (1♂ 1♀ NDSU). Ransom Co.: McLeod, Sandhills, 26-VIII-1972, D. Scott (1♂ lacking pygophore 1♀ with ♂ pygophore attached NDSU); 1mi. SE McLeod, 7-IX-1979, P. R. Larson (1♂ NDSU); 2mi. SW McLeod, T134N, R53W, Sec. 35, 14-VII-1992, D. A. Rider (1♂ DARC); 4mi. NW McLeod, T134N, R53W, Sec. 6, 18-VI-1992, D. A. Rider (1♂ DARC); Mirror Pool WMA, 46°32.348N, 97°17.475W, 27-VI-2012, D. A. Rider (1♂ DARC), 30-VIII-2007, D. A. Rider (2♀♀ DARC), 4-IX-2009, D. A. Rider (2♂♂ 4♀♀ DARC); 2mi. W Mirror Pool, 29-VII-1993, D. L. Cuthrell, at light (1♀ NDSU); 7mi. SE Sheldon, 5-IX-1979, E. U. Balsbaugh (2♂♂ 1♀ NDSU); SNG, A-Annex, 46.441N, 97.384W, 6-IX-2001, D. A. Rider (1♂ 1♀ DARC); 13-VII-1986, K. Larsen (1♂ MISU). Richland Co.: Ekre Grassland Preserve, T136N, R51W, Sec. 32, 28-VII-1993, D. L. Cuthrell (1♂ NDSU), 3-IX-2009, D. A. Rider (10 ♂♂ 16♀♀ DARC), 6-IX-1994, D. A. Rider (1♀ DARC), 24-IX-1994, K. Urlacher (2♀♀ NDSU), 25-IX-1994, T. Nowatzki (1♂ 2♀♀ NDSU); Mirror Pool, T135N, R52W, Sec. 8, 18-VII-1992, D. A. Rider (1♂ DARC), 28-VIII-1996, P. Anderson (1♂ NDSU), 24-IX-1994, K. Urlacher, at light (1♀ NDSU); 1.5mi. NE Mirror Pool, 18-VII-1993, D. L. Cuthrell (1♂ NDSU); SNG, Horsetrail Head, 46°31.409N, 97°12.194W, 27-VI-2012, D. A. Rider (1♀ 2012 THE GREAT LAKES ENTOMOLOGIST 367 DARC), 25-VIII-2009, D. A. Rider (9♂♂ 4♀♀ DARC), 4-IX-2009, D. A. Rider (5♂♂ 2♀♀ DARC), 9-IX-2008, D. A. Rider (6♂♂ 9♀♀ DARC); SNG, Oak Savannah, 17-IX-1995, P. Anderson (1♀ NDSU); SNG Orchid Site B, T134N, R53W, Sec. 35, 8-IX-1992, D. Cuthrell (1♂ NDSU); SNG, West I allotment, T135N, R52W, Sec. 10, 3-IX-2002, A. Joshi (2♂♂ NDSU); T136N, 61W, Sec. 32, 23-VII-1993, G. Fauske (1♂ NDSU); 6-VIII-1958, S. Wilson (2♂♂ NDSU). Slope Co.: Burning Coal Vein St. Pk., 1-VII-1965, L. Grochowski (1♀ NDSU), 10-VIII-1993, D. A. Rider (1♂ DARC), 12-VIII-1993, D. A. Rider (1♂ DARC). Homaemus bijugis Uhler, 1872 - Although this appears to be the most common scutellerid species occurring throughout the state, it appears that it has not actually been officially recorded from the state. The range given by McPherson (1982) for this species is Wisconsin and Illinois west and southwest to Saskatchewan and Alberta, Montana, Idaho, Wyoming, Colorado, and Ari- zona. Froeschner (1988d) listed it from Minnesota, Montana, South Dakota, and Saskatchewan, and Maw et al. (2000) recorded it from both Manitoba and Saskatchewan. Specimens Examined: NORTH DAKOTA [New State Record]: Barnes Co.: Katie Olson WMA, N of Valley City, 47°04′N, 98°04′W, 2-VII-2002, A. Joshi (1♀ NDSU); Rogers. Wesley’s Acres, 47°04′N, 98°10′W, 5-IX-2002, A. Joshi (1♀ NDSU); south of Valley City, 47°04′N, 98°04′W, 18-IX-2002, A. Joshi (1♀ NDSU). Billings Co.: Freeman’s Frontier nr. Medora, T140N, R102W, Sec. 26, 9-VIII- 1993, D. A. Rider (1♂ 3♀♀ DARC), 17-IX-1994, K. Urlacher (1♂ 2♀♀ NDSU); Theo. Roos. Mem. Prk, 25-VI-1964, R. L. Post, sweeping grass (1♂ NDSU), 29- VI-1965, L. Grochowski (1♀ NDSU). Cass Co.: Fargo (1♀ NDSU), VIII-1971, M. Knutson (1♀ NDSU), 17-IX-1948, R. L. Post (1♀ NDSU); Fargo, NDSU campus, 5-IX-2000, J. Kramer (1♂ NDSU), 6-IX-2001, D. A. Rider (1♂ DARC), 9-IX-2008, N. Noah (1♂ NDSU); 20-IX-1958, S. Wilson (1♂ NDSU), IX-1968 (1♀ NDSU), 15- X-1958 (1♂ NDSU), fall 1958 (1♀ NDSU). Dunn Co.: Killdeer Mts., 11-VI-1965, L. Grochowski, (2♂♂ NDSU), W. Kotchman (1♂ 1♀ NDSU); 2-VII-1976, Lago & Kurtz (1♀ NDSU). Golden Valley Co.: Beach, 10-VIII-1924, R. L. Webster (1♀ NDSU), 5-IX-1921, R. L. Webster (1? NDSU), 19-IX-1922 (1♀ NDSU). Hettinger Co.: 14mi. NE New England, T136N, R96W, Sec. 3, 17-IX-1994, K. Urlacher (3♂♂ 1♀ NDSU); Indian Creek Dam, T133N, R95W, Sec. 17, 8-X-1994, K. Urlacher (1♀ NDSU). McKenzie Co.: Th. Roos. M. Prk, 23-VI-1977, E. U. Balsbaugh (1♂ NDSU), 20-VII-1965, L. Grochowski (1♀ NDSU), 4-VIII-1965, L. Grochowski (1♀ NDSU); 2-VIII-1977, L. A. Schutz & J. W. Smith (1♀ NDSU). McLean Co.: 24- III-1957, P. Bergen (1♀ NDSU), 10-VII-1957, P. Bergen (1♀ NDSU). Morton Co.: Mandan, 11-VIII-1922, R. L. Webster (1♂ NDSU), 12-VIII-1922, R. L. Webster (1♀ NDSU), 18-VIII-1922, R. L. Webster (1♀ NDSU); 19-VIII-1958, R. L. Post (3♂♂ 1♀ NDSU). Oliver Co.: 2mi. E Hensler, 20-VII-1978, L. A. Schutz & J. W. Smith (1♂ NDSU). Ransom Co.: 1.5mi. W McLeod, T134N, R53W, Sec. 22, 4-VI-1992, D. A. Rider (1♂ DARC); 1.5mi. SE McLeod, T134N, R52W, SW 31, 11-VIII-1992, D. Cuthrell (1♀ NDSU); 2mi. SW McLeod, T134N, R53W, Sec. 35, 14-VII-1992, D. A. Rider (1♂ DARC), 17-VII-1992, D. A. Rider (4♀♀ DARC); 4mi. NW McLeod, T134N, R53W, Sec. 6, 18-VI-1992, D. A. Rider (3♂♂ 2♀♀ DARC); SNG, A-Annex, 46.441N, 97.384W, 6-IX-2001, D. A. Rider (4♂♂ 3♀♀ DARC); SNG, Orchid site B, T134N, R53W 35, 5-VIII-1992, D. Cuthrell (1♀ NDSU); SNG, Orchid site D, T134N, R54W, SE 12, 3-IX-1992, D. Cuthrell (1♂ 1♀ NDSU); SNG, 9-IX-2008, K. Ghising (1♂ NDSU); 1-VII-1964, R. L. Post (1♂ 4♀♀ NDSU), 6-VIII-1958, S. Wilson (1♂ 1♀ NDSU). Richland Co.: Ekre Grassland Preserve, T136N, R51W, Sec. 32, 6-IX-1994, M. Ellefson (1♂ NDSU), T. Nowatzki (1♂ NDSU), D. A. Rider (2♀♀ DARC), W. G. Schroeder (2♂♂ 1♀ NDSU); McLeod, T133N, R52W, Sec. 6, 23-VIII-2000, J. Maxwell (1♂ NDSU), 18-IX-1979, E. U. Bals- baugh (3♀♀ NDSU); 1.5mi. SE McLeod, T134N, R52W, Sec. 31, 8-IX-1992, D. Cuthrell (1♀ NDSU); Mirror Pool, T135N, R52W, Sec. 8, 27-VI-1992, D. A. Rider (4♂♂ 1♀ DARC), 2-VII-1979, J. W. Hoganson (1♂ 1♀ NDSU), 4-IX-1998, D. A. Rider (1♂ 3♀♀ DARC), 5-IX-1994, T. Nowatzki (1♀ NDSU), 15-IX-1994, 368 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

K. Urlacher (2♀♀ NDSU), 18-IX-1994, D. A. Rider (2♀♀ DARC), 25-IX-1994, K. Urlacher, UV light (2♂♂ 1♀ NDSU); 1.5mi. NE Mirror Pool, 15-VI-1994, D. A. Rider (3♀♀ DARC); 1mi. NE Mirror Pool, 25-VI-1993, D. L. Cuthell (1♂ 1♀ NDSU), 17-VII-1993, D. L. Cuthrell (1♀ NDSU), 28-VII-1993, D. L. Cuthrell (1♀ NDSU); 2mi. NE Mirror Pool, 28-VI-1993, G. Fauske (1♀ NDSU); SNG, Horsetrail Head, 46°31.409N, 97°12.194W, 27-VI-2012, D. A. Rider (8♂♂ 5♀ DARC), 20-VII-2012, D. A. Rider (5♂♂ 3♀♀ DARC), 25-VIII-2009, D. A. Rider (1♂ DARC), 9-IX-2008, D. A. Rider (1♂ 2♀♀ DARC), 25-X-2007, D. A. Rider (2♀♀ DARC); SNG, Oak Savannah area, T135N, R52W, Sec. 10, 18-VI-2000, D. A. Rider (3♂♂ 2♀♀ DARC), 28-VI-1995, G. Fauske (2♂♂ 5♀♀ NDSU), 17- VII-1993, D. Cuthrell (1♂ NDSU), 26-IX-1995, G. Fauske (3♂♂ 4♀♀ NDSU); SNG, I allotment, T235N, R52W, Sec. 10, 3-IX-2002, A. Joshi (1♂ 2♀♀ NDSU); Walcott Dunes 28-VIII-1971, T. L. McCabe (1♀ NDSU); T135N, 52W, Sec. 8, 12-IX-1992, J. A. Nelson (2♂♂ NDSU); T136N 61W Sec. 32, 23-VII-1993, G. Fauske (1♂ NDSU); 1-VII-1964, R. J. Sauer (1♂ NDSU), 19-VII-1965, R. L. Post (1♀ NDSU), 4-VIII-1958, S. Wilson (1♀ NDSU), 6-VIII-1958, S. Wilson (1♂ 1♀ NDSU), 13-IX-1968, R. Johnson (1♀ NDSU). Slope Co.: Burning Coal Vein St. Pk., 28-VI-1982, R. J. Dregseth & T. McCabe (1♂ NDSU), 1-VII-1965, L. Grochowski (2♂♂ 1♀ NDSU), 1-VIII-1972, T. L. McCabe & R. J. Dregseth (1♀ NDSU), 10-VIII-1993, D. A. Rider (1♂ DARC); Chalky Buttes, 7-VI-1965, L. Grochowski (1♂ NDSU); T135N, R102W, Sec. 27, west edge, 19-VII-1978, L. A. Schutz & J. W. Smith (1♀ NDSU); 11-VII-1986, K. Larsen (1♀ MISU), 8-VIII-1976 (1♀ NDSU), 20-VIII-1955, V. Goodfellow (1♀ NDSU), 20-VIII-1958, R. L. Post (1♀ NDSU). Ward Co.: SE Minot, behind Menards, 17-IX-1999, G. Hanley (5♂♂ MISU).

Tetyra Fabricius, 1803 The genus Tetyra contains six species, most of which are neotropical in distribution. Two of the neotropical species occur in the United States only in the southern states. For example, T. antillarum Kirkaldy occurs in Florida southward; T. robusta Uhler occurs in Arizona, Nevada, and Utah, southward into Mexico. There is only one species that is more widely distributed in the eastern United States. Tetyra bipunctata (Herrich-Schäffer) has not yet been recorded from North Dakota, but Froeschner (1988d) listed it from Minnesota, and the NDSU collection contains a specimen from northwestern Minnesota: Norman Co.: Gary Pines, 17-X-2000, R. Halland (1♀ NDSU). Norman County is on the border with North Dakota, so it is likely this species will eventually be collected in North Dakota. It feeds on and has been recorded as a pest of various species of pine (Gilbert et al. 1967).

THYREOCORIDAE Amyot and Serville, 1843 Members of this family have in the past been called negro bugs (because of their general black color). It has at times been considered to be a subfamily of a very inclusive Cydnidae (Dolling 1981), but most present-day workers treat it as a full family. Although it is becoming dated, the single best publication to help identify thyreocorids is that of McAtee and Malloch (1933). Some caution is needed as some names have changed, and there have been a number of new species described since 1933, but it is impressive how useful this paper remains. Identification of thyreocorids can be tedious, partially because of the small size of many of its species; but also some of the important characters involve examining the location of the spiracles and associated trichobothria. Fortunately, our species are relatively easy to identify. Only three genera are known to occur in our area. Thyreocorids tend to live at or near the ground, but they are often encountered higher up on host plants, especially members of the Apiaceae. 2012 THE GREAT LAKES ENTOMOLOGIST 369

Key to genera of North Dakota Thyreocoridae 1 Anterolateral pronotal margins and costal margins of coria with long slender bristles...... Cydnoides – Anterolateral pronotal margins and costal margins of coria lacking long slender bristles...... 2 2(1) Coloration completely black, no pale markings...... 3 – Coloration black with at least some pale markings along basal costal margins of coria...... Corimelaena (part) 3(2) Spiracles nearer to trichobothria than to lateral margins of abdominal sternites; costal margin bordered within by definite groove...... Galgupha – Spiracles nearer to lateral margins of abdominal sternites than to trichobothria; costal margin not bordered within by definite groove...... Corimelaena (part)

Corimelaena White Although there is some overlap, North American species of Corimelaena tend to be smaller (less than 4.8mm in length) than species of Galgupha (most are greater than 4.5mm in length). Nearly all North American species of Corimelae- na have pale markings on the basal costal margin of each corium, the exception being C. nigra. In contrast, nearly all North American species of Galgupha lack pale markings on the corium (it should be noted that many neotropical species of Galgupha have pale markings on the coria, several of which may enter the southern parts of North America). Although only two Corimelaena species are positively known from North Dakota, there are three more species that may eventually be collected here. Froeschner (1988e) listed C. alpina McAtee from Minnesota, even though McPherson (1982) indicated that the Minnesota record came from an unpublished manuscript by R. Sailer. Corimelaena nigra Dal- las has been recorded from both north and south of North Dakota (Froeschner 1988e, Maw et al. 2000), but I have not examined any specimens from our region. Corimelaena extensa Uhler appears to be a western species, but was listed by Froeschner (1988e) from Montana and South Dakota, so it may eventually be found in North Dakota. The following key includes all five species in anticipa- tion of eventually finding any of the above mentioned species.

Key to species of Corimelaena that may occur in North Dakota 1 Coloration completely black, lacking pale markings on coria [not yet recorded from North Dakota]...... nigra Dallas – Coloration black with pale markings along basal costal margin of each corium...... 2 2(1) Pale markings along basal lateral margin of each corium confined to exocorium, not extending onto mesocorium...... lateralis (Fabricius) – Pale markings along basal lateral margin of each corium extending onto mesocorium...... 3 3(2) Spiracles of segments three to six below the lateral carina; apex of corium obtusely rounded...... 4 – Spiracles of segments three to six in the lateral carina; apex of corium pointed, not rounded [not yet recorded from North Dakota]...... extensa Uhler 4(3) Hind tibia without posterodorsal bristles; sides of head in front of eyes slightly emarginate; head from lateral view only slightly arched [not yet recorded from North Dakota]...... alpina (McAtee and Malloch) 370 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

– Hind tibia with one to three short posterodorsal bristles; sides of head in front of eyes distinctly emarginate, especially in males; head from lateral view strongly arched...... pulicaria (Germar) Corimelaena (Corimelaena) lateralis (Fabricius, 1803) - This species is relatively easy to identify; it is the only thyreocorid species in our area having the pale lateral margin of the coria confined to the exocorium, not spilling onto the mesocorium basally. It also tends to be slightly larger than our other common species, C. pulicaria. Apparently, there have been no specific localities publishied for North Dakota. Neither Uhler (1876) nor Hussey (1922) cited this species from North Dakota. McAtee and Malloch (1933) provided the following range: “New York to North Dakota and south to Florida and Texas.” Possibly based on the range given by McAtee and Malloch, Froeschner (1988d) listed this species from North Dakota. This species has been reared in the laboratory (McPherson 1971a); he (1972) also studied its life history and described the immature stages. Specimens Examined: NORTH DAKOTA [New State Record]: Rich- land Co.: Mirror Pool, T135N, R52W, NE 8, 3-VI-1994, D. A. Rider (3♂♂ 1♀ DARC), 27-VI-1992, D. A. Rider (1♀ DARC, 1♂ NDSU). Corimelaena (Corimelaena) pulicaria (Germar, 1839) - In this species, the pale lateral margin of the corium spills onto the mesocorium basally. There are two other species that may have this color pattern that could occur in North Dakota. Corimelaena extensa has a different arrangement of the spiracles (see the key above) and the apex of the corium is more sharply pointed than in this species; C. alpina lacks several bristles on the hind tibia that are present in this species (also see key above). This species tends to be slightly smaller than our other common species, C. lateralis. McPherson (1982) gave the range for this species as Québec, Ontario, and Massachusetts west to British Columbia and Oregon, and south to Florida, Arkansas, Texas and Guatemala. Uhler (1878) cited this species simply from ‘Dakota.’ Hussey (1922), however, provided three specific localities for North Dakota:Cass Co.: Fargo, T. H. Hubbell; Eddy Co.: near Sheyenne River, about 3 miles south of Warwick, on vegetation in an open woods, August 8, 1920, T. H. Hubbell; and Ramsey Co.: Devils Lake, August 14, 1920, on Ambrosia, T. H. Hubbell. Lugger (1900), in Minnesota, provided a colorful description of the unfortunate finding of these bugs in a plate of berries. Specimens Examined: NORTH DAKOTA: Barnes Co.: 1984, D. A. Mundal, Euphorbia esula (1♂ 2♀♀ NDSU). Billings Co.: Medora, 47°02′N, 103°36′W, 24-VI-2002, A. Joshi (2♀♀ NDSU); Th. Roos. M. Pk., 10-VI-1965, W. Kotchman (1♀ NDSU). Burleigh Co.: Bismarck, 24-VII-2007, D. Nelson, Lindgren trap (1♀ NDSU). Cass Co.: Fargo (1♂ 4♀♀ NDSU), 11-V-1940 (1♂ NDSU), 13-V-1939, W. Southam (1♀ NDSU), 20-V-1923, W. Baker (1♂ NDSU), 26-VII-1922, O. A. Stevens, Sonchus arvensis (3♀♀ NDSU), 17-IX- 1948, R. L. Post (1♀ NDSU); NE Fargo, 46.759N, 96.765W, 31-VII-2012, D. A. Rider (2♂♂ 1♀); N Fargo, Riverwood Pk., 46.945N, 96.802W, 16-VII-2012, D. A. Rider (1♀ DARC); Fargo, NDSU Campus, 21-V-2012, D. A. Rider (6♀♀ DARC), 1-VI-2011, D. A. Rider & G. Fauske, lindgren trap with Ips lure (1♀ NDSU), 14-VI-2011, D. A. Rider & G. Fauske, lindgren trap with Sirex lure (1♂ NDSU), 23-VIII-2002, F. Antwi (1♀ NDSU); S Fargo, 76 Ave. S, Chrisan Edition, 46.766N, 96.780W, 5-VI-2012, D. A. Rider (1♂ 2♀♀ DARC); Wild Rice R. at Red R., 46.758N, 96.787W, 1-VIII-2012, D. A. Rider (3♂♂ 2♀♀ DARC), 13-VIII-2012, D. A. Rider (2♂♂ 1♀ DARC); Wild Rice R. at University Dr., 46.759N, 96.797W, 27-VII-2012 (15♂♂ 9♀♀ DARC), 1-VIII-2012, D. A. Rider (2♂♂ DARC); 3-VIII-1968, J. Johnson (1♂ NDSU). Dunn Co.: Killdeer Mtns., 3-VIII-1965, W. Kotchman (1♂ 1♀ NDSU); 9-VIII-1976, Lago & Kurtz (1♀ NDSU). Golden Valley Co.: 1984, D. A. Mundal, Euphorbia esula (1♂ 2012 THE GREAT LAKES ENTOMOLOGIST 371 3♀♀ NDSU). Grand Forks Co.: Goose R., 1-VIII-1962, D. G. Aarhus (2♂♂ 1♀ NDSU); Inkster Spring, 20-X-1962, R. Poole (1♀ NDSU); Northwood, 25-V-1956, R. L. Post (3♂♂ NDSU), 27-V-1956, R. L. Post (1♂ 1♀ NDSU); Oakville Prairie, Sec. 16, 47°53.403N, 97°18.855W, 14-VIII-2009, D. A. Rider (1♂ DARC); 28-VIII-1958, R. C. & R. L. Post (1♀ NDSU). Hettinger Co.: 4mi. N Regent, T135N, R94W, Sec. 19, 18-VIII-1996, K. Urlacher, Fagopyrum esculintum (3♂♂ 3♀♀ NDSU); 5mi. N Regent, T135N, R94W, Sec. 18, 24-VII- 1995, K. Urlacher, pitfall trap in Brassica juncea (3♀♀ NDSU). McKenzie Co.: Th. Roos. M. Park, 22-VI-1965, L. Grochowski (1♀ NDSU), 24-VI-1964, R. L. Post (2♀♀ NDSU). Pembina Co.: 2-VII-1961, R. L. Post (1♂ NDSU). Ransom Co.: Lisbon, 1-VI-1983, J. Julian, on leafy spurge (1♀ NDSU), 11- VI-1982, J. Julian, on leafy spurge (1♂ NDSU); 7mi. SE Sheldon, 3-IX-1979, E. U. Balsbaugh (2♂♂ NDSU), 5-IX-1979, E. U. Balsbaugh (1♂ NDSU); SNG, A-Annex, T134N, R53W, Sec. 8, 26-V-1992, D. A. Rider (1♂ DARC). Richland Co.: SNG, Horse Trail Head, 46.523N, 97.201W, 28-V-2010, D. A. Rider (1♂ 9♀♀ DARC, 1♂ 3♀♀ NDSU), 20-VII-2012, D. A. Rider (1♀ DARC); SNG, Oak Savannah, T135N, R52W, Sec. 10NE, 23-VII-1996, L. Pederson (1♀ NDSU); 6-V-1962, R. D. Frye, grass (1♀ NDSU). Rolette Co.: 4-VI- 1968, Kotchman (3♂♂ 1♀ NDSU), 4-VI-1968, Kotchman, P. tremuloides (1♀ NDSU), 5-VI-1968, R. Katayama (1♂ NDSU). Slope Co.: 22-VI-1976, Lago & Kurtz (1♀ NDSU). Stark Co.: Dickinson, Patterson Lake, 9-VIII-2011, K. Marmon, lindgren trap with AP + GE lure (1♀ NDSU), 23-VIII-2011, K. Marmon, lindgren trap with Ips lure (4♀♀ NDSU), 23-VIII-2011, K. Marmon, lindgren trap with AP + GE lure (2♀♀ NDSU), 23-VIII-2011, K. Marmon, lindgren trap with Chalcogram (1♀ NDSU). Towner Co.: 6-VI-1968, R. W. Katayama (1♀ NDSU), 6-VI-1968, Kotchman (1♂ 1♀ NDSU). Ward Co.: Minot, 14-V-1941, L. E. Storhaug (1♂ MISU). Williams Co.: 1984, D. A. Mundal, Euphorbia esula (2♀♀ NDSU).

Cydnoides Malloch, 1919 To date, no species of Cydnoides has been recorded from North Dakota; I have not examined any specimens from this area either. There are, however, three species that may eventually be collected in North Dakota. One paratype of the subspecies C. ciliatus orientis McAtee and Malloch (1933) is from Scott Co., Minnesota, which is just south of Minneapolis. Cydnoides renormatus (Uhler) appears to be primarily a desert southwestern species, but Hart (1919) recorded one specimen from northern Illinois, and according to McPherson (1982), Sailer in an unpublished manuscript mentioned an additional specimen from Minnesota. Harris (1943) recorded C. albipennis (Say) from South Dakota. I have provided a key to the identification of Cydnoides species that might occur in North Dakota.

Key to species of Cydnoides that may occur in North Dakota (extracted from McAtee and Malloch, 1933) 1 Hind tibia without a carinate line on posterior surface; exposed corium about as wide at apex as at base (subgenus Sayocoris)....albipennis (Say) – Hind tibia with a definite carinate line on posterior surface; exposed corium not nearly as wide as apex as at base (subgenus Cydnoides)...... 2 2(1) Corium entirely fuscous to black, no part whitish; hind tibia with at least five posterodorsal bristles...... ciliatus Uhler – Corium partly white or cream-colored; hind tibia with not more than four posterodorsal bristles...... renormatus Uhler 372 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

Galgupha Amyot and Serville, 1843 Galgupha is an extremely diverse genus, with many species, especially in the neotropics. As mentioned previously, all our species are essentially solid black in color. Some of the southern U.S. species and many of the neotropical species, however, have pale markings not only on the corium, but sometimes also on the scutellum. Our species tend to be slightly larger than species of Corimelaena. Although the pale markings on the corium will usually separate our common species of Corimelaena from this genus, one may need to examine the position of the spiracles in relation to the trichobothria, or become familiar with the grooves and/or ridges along the margin of the corium to separate some species (see key to genera above). Three species of Galgupha are known to occur in North Dakota; a fourth species (G. loboprostethia Sailer) may eventually be found here (in fact, see discussion under G. ovalis; possibly it has been found here). Froeschner (1988e) listed G. loboprostethia from South Dakota; it can be recognized by the expanded prostethium.

Key to species of Galgupha that may occur in North Dakota 1 Lateral area of metapleuron distinctly punctate laterad of evaporative areas (subgenus Nothocoris McAtee and Malloch)...... nitiduloides (Wolff) – Lateral area of metapleuron impunctate laterad of evaporative areas (subgenus Galgupha Amyot and Serville)...... 2 2(1) Anterior margin of prostethium produced to form explanate lobe, granulated portion as wide as base of second rostral segment [not yet recorded from North Dakota]...... loboprostethia Sailer – Anterior margin of prostethium not lobed, granulated portion relatively narrow...... 3 3(2) Anterodorsal series of spines on each fore tibia with distal two spines distinctly weaker than preceding spines, or lacking; corium without distinct ridge inside costal groove, ridge represented only by series of punctures distally; punctation on discal portions of pronotum and scutellum mostly obsolete...... atra Amyot and Serville – Distal two spines of anterodorsal series on fore tibiae subequal in size with preceding spines; corium with distinct ridge inside costal groove; punctation on discal portions of pronotum and scutellum more evident ...... ovalis Hussey Galgupha (Galgupha) atra Amyot and Serville, 1843 - This species lacks the punctures laterad of each evaporative area; separation of this species from G. ovalis requires a fairly detailed examination of the spines on the fore tibiae, and/or the ridges on the coria (see key above). McPherson (1982) gave the distribution of this species as Québec, Maine, and Washington south to Florida, Texas, Arizona, and Mexico. Froeschner (1988e) listed this species from Manitoba, Montana, North Dakota, and South Dakota. Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. Specimen examined: NORTH DAKOTA: Billings Co.: Roosevelt Natl. Pk., Medora, 24-VI-2002, F. Antwi (2♀♀ NDSU). Cass Co.: Fargo, 22-V-1923, W. Baker (1♀ NDSU). Ransom Co.: 2mi. SE McLeod, T134N, R53W, Sec. 35 SW 1/4, 8-VII-1992, D. Cuthrell & T. Madson, Hg light (1♀ NDSU); 7mi. SE Sheldon, 5-IX-1974, J. W. Smith (1♀ NDSU). Richland Co.: Walcott Dunes, T. L. McCabe (1♀ NDSU). Rolette Co.: Turtle Mtns., 8-VII-1964, R. J. Sauer (1♀ NDSU). Starck Co.: Dickinson, 8-VII-1941, H. S. Telford (1♀ NDSU). Galgupha (Galgupha) ovalis Hussey, 1925 - This species lacks the punctures laterad of each evaporative area; to separate it from G. atra, please 2012 THE GREAT LAKES ENTOMOLOGIST 373 see the characters in the key (above). McPherson (1982) gave the distribution of this species as Massachusetts west to Montana, and south to Florida, Arizona, Texas, and Guatemala. Neither Uhler (1876) nor Hussey (1922) recorded this species from North Dakota. Apparently, it has not previously been recorded from North Dakota. Froeschner (1988e) listed this species from Montana, South Dakota, and many states to the east and south of North Dakota. Interestingly, one of the female specimens in the series collected in Grant Co. on May 18, 1964, has the prostethium expanded similarly as seen in specimens of G. loboprostethia, but it is identical in all other respects with the remaining specimens in the series. At this time, it seems prudent to consider this specimen as an aberrant individual of G. ovalis. It is also interesting to note that all specimens of this species collected so far from North Dakota come from either Grant County or Logan County, both of which are in the south central area of North Dakota. Specimens Examined: NORTH DAKOTA [New State Record]: Grant Co.: Grant Co.: Heart Butte Dam, 18-V-1964, R. L. Post (2♂♂ 3♀♀ NDSU); 18- V-1964 (6♂♂ 3♀♀ NDSU). Logan Co.: 16-VI-1967, W. Kotchman (1♂ 1♀ NDSU). Galgupha (Nothocoris) nitiduloides (Wolff, 1802) - This is the easiest species to recognize in our area. The punctures laterad of each evaporatorium are diagnostic, although be aware that in greasy specimens, it is almost impossible to see these punctures. Also, the number of punctures is not great, and the area of punctures is not extensive, but rather it is more in the form of a narrow band paralleling the margin of the evaporative area. Hussey (1922) stated that “Van Duzee (1917, p. 14, No. 28) lists this species as occurring in North Dakota. It does not occur in Mr. Hubbel’s collection.” Froeschner (1988e) listed this species from Minnesota, Montana, and North Dakota, and Maw et al. (2000) recorded this species from both Manitoba and Saskatchewan. Specimens Examined: NORTH DAKOTA: Barnes Co.: 27-VI-1958 (1♀ NDSU). Burleigh Co.: Bismarck, landfill #2, 6-VII-2011, C. Larson, Lindgren trap (1♀ NDSU). Cass Co.: Fargo, 10-VII-1924, A. Oderkirk, swept from flax (1♂ NDSU). Grand Forks Co.: Inkster Spring, 13-VII-1964, R. J. Sauer (1♂ NDSU), 17-VIII-1961 (1♂ NDSU). Oliver Co.: 10mi. SE Kensler, T142N, R81W, Sec. 7, 20-VII-1995, K. Urlacher, pitfall trap in Fagopyrum esculintum (1♀ NDSU). Ransom Co.: SNG, A Annex, T134N, R53W, Sec. 8, 14-VII-1992, D. A. Rider (1♀ DARC), 17-VII-1992, D. L. Cuthrell (1♀ NDSU); SNG, 1.5mi. SE McLeod, T134N, R52W, SW31, 7-9-VIII-1992, D. L. Cuthrell (1♂ NDSU); SNG, 2mi. SE McLeod, 14-VII-1992, D. A. Rider (1♂ DARC); SNG, 4mi. NW McLeod, 25-VI- 1992, D. A. Rider (1♂ DARC); 1mi. NE Mirror Pool, 17-VII-1993, D. A. Rider (4♂♂ 2♀♀ DARC). Richland Co.: Mirror Pool, T135N, R52W, Sec. 8, 18-VII- 1992, D. A. Rider (1♀ DARC), 14-21-IX-1994, W. G. Schroeder (1♂ NDSU); 1mi. E Mirror Pool, 28-VII-1993, D. A. Rider (1♀ DARC); 1mi. NE Mirror Pool, 17- VII-1993, D. A. Rider (2♂♂ 1♀ DARC), 27-VII-1993, D. L. Cuthrell (1♂ NDSU); SNG, Horse Trail Head nr. Oak Savannah46.52N, 96.21W, 20-VII-2012, D. A. Rider (4♂♂ DARC), 25-X-2007, D. A. Rider (1♂ DARC); Wahpeton, 8-XI-1964, L. Fomes & R. L. Post, in sod by wheat (1♀ NDSU); 10mi. W Walcott, 3-V-1978, E. U. Balsbaugh (1♂ 2♀♀ NDSU); SNG, T135N, R52W, Sec. 30, 5-VII-2000, J. Maxwell (1♂ NDSU).

Acknowledgments I would like to thank Jeff Vaughn, University of North Dakota (UNDG), and Guy Hanley, Minot State University (MISU) for allowing me to study the collections in their care. I also thank my technician, Jerry Fauske, past graduate students and student workers (too numerous to name) all of whom collected many of the specimens cited in this study. 374 THE GREAT LAKES ENTOMOLOGIST Vol. 45, Nos. 3 - 4

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