Dr. Richard Clopton

Professor of Biology

Dr. Richard Clopton

Professor of Biology

Arts & Sciences
Hoyt 312


Office/Lab Hours:
MW / 8 am – 9:15 am & 12:30 pm – 1:45 pm
TR / 8 am – 9:15 am


Fall 2018

Coll 101-000C / College Success (Science/Math)
Biol 102-000A / Introductory Zoology
Biol 311-000A / Evolution & Systematic Biology

Spring 2019



Laboratory for Gregarine Parasitology Website ( Hotel Intestine)


I teach courses in evolution, systematics, and organismal biology. My research interests include evolution, systematics, biodiversity and natural history of invertebrates and their parasites. I am particularly interested in understanding the forces underlying speciation in hyperdiverse parasitic lineages. My research program is supported through consistent competitive grant support from the National Science Foundation and we have recently finished projects on the gregarine parasites of insects in the Nebraska Sandhills, the Texas “Big Thicket”, and Belize, Central America (photo above). I have recently completed a long-term molecular phylogeny project to resolving major trends in gregarine evolution and establishing the first evolutionary systematic arrangement of the group. My current research is focused on the distribution and biodiversity of North American cockroaches and their gregarine parasites.

My research program incorporates undergraduate research students as research colleagues: they conduct original science, present their findings at regional and national meetings, and publish their results in peer-reviewed scientific journals. Science is a discipline learned by working with other scientists and I try to maintain a vigorous, enthusiastic, and productive laboratory. I am always looking for students who are interested and motivated to become scientists – feel free to stop by the laboratory or visit my research website: http:science.peru.edu/gregarina


  • R. E. Clopton. 2011. Redescription of Protomagalhaensia granulosae Peregrine, 1970 (Apicomplexa: Eugregarinida: Blabericolidae) parasitizing the Discoid Cockroach, Blaberus discoidalis (Dictyoptera: Blaberidae). Comparative Parasitology 78 (In press).
  • Cielocha, J. J., T. J. Cook, R. E. Clopton. 2011. Host Utilization and Distribution of Nubenocephalid Gregarines (Eugregarinorida:  Actinocephalidae) Parasitizing Argiaspp. (Odonata: Zygoptera) in the Central United States. Comparative Parasitology 78 (In review).
  • Cook, T. J., R. E. Clopton, and J. L. Cook. 2011. Neoasterophora megasylvaticus n. gen., n. sp. (Apicomplexa: Eugregarinida: Actinocephalidae: Pileocephalinae) Parasitizing Cheumatopsyche analis (Trichoptera: Hydropsychidae) in the Texas Big Thicket. Comparative Parasitology 78 (In Press).
  • Clopton, R. E., T. J. Cook, and J. J. Cielocha. 2010. Nubenocephalus nickoli n. sp. and Nubenocephalus xunantunichensis n. sp. (Apicomplexa: Eugregarinida: Actinocephalidae) Parasitizing Damselflies (Odonata: Zygoptera) in Belize, Central America. Comparative Parasitology 77 (In press).
  • Clopton, R. E.. 2010. Protomagalhaensia cerastes n. sp. (Apicomplexa: Eugregarinida: Blabericolidae) Parasitizing the Pallid Cockroach, Phoetalia pallida(Dictyoptera: Blaberidae). Comparative Parasitology 77 (In press).
  • Clopton, R. E.. 2009. Phylogenetic Relationships, Evolution, and Systematic Revision of the Septate Gregarines (Apicomplexa: Eugregarinorida: Septatorina). Comparative Parasitology 76: 167-190.
  • Clopton, R. E., T. J. Cook, and J. L. Cook. 2008. Gregarina tropica n. sp. (Apicomplexa: Eugregarinorida: Gregarinicae: Gregarinidae) Parasitizing the Brown-winged Earwig, Vostox brunneipennis (Dermaptera: Labiidae), in the Texas Big Thicket. Comparative Parasitology 75:215-227.
  • Clopton, R. E., T. J. Cook, and J. L. Cook. 2008. Trichurispora wellgundis n. g., n. sp. (Apicomplexa: Eugregarinida: Hirmocystidae) parasitizing adult Water Scavenger Beetles, Tropisternus collaris (Coleoptera: Hydrophilidae) in the Texas Big Thicket. Comparative Parasitology 75: 82-91.
  • Realzola, E., J. L. Cook, T. J. Cook, and R. E. Clopton. 2007. Composition of gyrinid aggregations in the East Texas Primitive Big Thicket.  Coleopterist’s Bulletin 61:495-502.
  • Hays, J. J., R. E. Clopton, T. J. Cook and J. L. Cook. 2007. Revision of the GenusNubenocephalus and Description of Nubenocephalus secundus N. Sp. (Apicomplexa: Actinocephalidae) Parasitizing Adults of Argia sedula (Odonata: Zygoptera: Coenagrionidae) in the Primitive Texas Big Thicket, U.S.A. Comparative Parasitology. 74: 286-293.
  • Clopton, R.E., T. J. Cook and J. L. Cook. 2007. Revision of GeneiorhynchusSchneider, 1875 (Apicomplexa: Eugregarinida: Actinocephalidae: Acanthosporinae) with Recognition of 4 New Species of Geneiorhynchus and Description ofGeneiorhynchus manifestus N. Sp. Parasitizing Naiads of the Green Darner, Anax junius (Odonata: Aeshnidae) in the Texas Big Thicket. Comparative Parasitology 74: 273-285.
  • Clopton, R. E., and J. J. Hays. 2006. Revision of the Genus Protomagalhaensia and Description of Protomagalhaensia wolfi n. comb. (Apicomplexa: Eugregarinida: Hirmocystidae) and Leidyana haasi n. comb. (Apicomplexa: Eugregarinida: Leidyanidae) Parasitizing the Lobster Cockroach, Nauphoeta cinerea (Dictyoptera: Blaberidae) Comparative Parasitology 73: 137-156.
  • Clopton, R. E. 2006. Two New Species of Xiphocephaus in Eleodes tricostata andEleodes fusiformis (Coleoptera: Tenebrionidae: Eleodini) from The Sandhills of Western Nebraska. Journal of Parasitology  92: 569-577.
  • Welsh, D., R. E. Clopton, and L. Parris. 2006. Differential Temperature Acclimatization Responses in the Membrane Phospholipids of Posthodiplostomum minimum and its Second Intermediate Host, Lepomis macrochirus. Journal of Parasitology 92: 764-769.
  • Snyder, S. D., and R. E. Clopton. 2005. New Methods for the Collection and Preservation of Spirorchiid and Polystomatid Trematodes from Turtles. Comparative Parasitology 72: 102-107.
  • Hays, J., R. E. Clopton, D. L. Cappaert, and D. R. Smitley. 2004. Revision of the genus Stictospora and description of Stictospora villani, n. sp. (Apicomplexa: Eugregarinida: Actinocephalidae) from the Japanese beetle Popillia japonica(Coleoptera: Scarabaeidae) in Michigan, USA. Journal of Parasitology 90: 1450-1456.
  • Clopton, R. E.  2004. Standard nomenclature and metrics of plane shapes for use in gregarine taxonomy. Comparative Parasitology 71: 130-140.
  • Clopton, R. E.  2004. Calyxocephalus karyopera g. nov., sp. nov. (Eugregarinorida: Actinocephalidae: Actinocephalinae) from the ebony jewelwing damselfly Calopteryx maculata (Zygoptera: Calopterygidae) in southeast Nebraska: implications for mechanical prey-vector stabilization of exogenous gregarine development. Comparative Parasitology 71: 141-153.
  • Clopton, R. E., T. J. Cook, and J. L. Cook.  2004.  Naiadocystis phykoterion n. gen., n. sp. (Apicomplexa: Eugregarinida: Hirmocystidae) from the mexican pygmy grasshopper Paratettix mexicanus (Orthoptera: Tetrigidae) in the Texas Big Thicket with recognition of three previously described species of Naiadocystis. Journal of Parasitology 90: 301-307.
  • Clopton, R. E.  2003.  Comparative Parasitology policies and instructions for authors. Comparative Parasitology 70: 218-225.
  • Smith A., and R. E. Clopton.  2003.  Efficacy of oral metronidazole and potassium sorbate against two gregarine parasites, Protomagalhaensia granulosae andGregarina cubensis (Apicomplexa: Eugregarinida) infecting the Death’s Head Cockroach, Blaberus discoidalis. Comparative Parasitology 70: 196-199.
  • Leander, B. S., R. E. Clopton, and P. J. Keeling.  2003.  Phylogeny of gregarines (Apicomplexa) as inferred from small-subunit rDNA and beta-tubulin. International Journal of Systematic and Evolutionary Microbiology 53: 345-354.
  • Clopton, R. E.  2003.  Comparative Parasitology policies and instructions for authors. Comparative Parasitology 69: 223-230.
  • Clopton, R. E., and C. M. Nolte.  2002.  Clitellocephalus americanus n. gen., n. sp. (Apicomplexa: Eugregarinida: Gregarinidae) from Cratacanthus dubius (Coleoptera: Carabidae: Harpalinae) in the Nebraska Sandhills and Clitellocephalus ophoni n. comb. (Apicomplexa: Eugregarinida: Gregarinidae) from Ophonus pubescens(Coleoptera: Carabidae: Harpalinae) in Sète, France. Journal of Parasitology 88: 750-757.
  • Clopton, R. E., and A. Smith.  2002.  Efficacy of oral sulfadimethoxine against two gregarine parasites, Protomagalhaensia granulosae and Gregarina cubensis(Apicomplexa: Eugregarinida) infecting the Death’s Head Cockroach, Blaberus discoidalis. Journal of Parasitology 88: 786-789.
  • Clopton, R. E.  2002.  Phylum Apicomplexa Levine, 1970: Order Eugregarinorida Léger, 1900.  Pages 205-288 in:  Illustrated Guide to the Protozoa, 2nd edition, J. J. Lee, G. Leedale, D. Patterson, and P. C. Bradbury, eds.  Society of Protozoologists, Lawrence, Kansas.
  • Cook, T. J., J. Janovy, Jr., and R. E. Clopton.  2001. Epimerite-host epithelium relationships among eugregarines parasitizing the damselflies Enallagma civile andIschnura verticalis.  Journal of Parasitology 87: 988-996.
  • Clopton, R. E.  2000.  Stylocephalus occidentalis n. sp.(Apicomplexa: Eugregarinida: Stylocephalidae) from Trimytis pruinosa (Coleoptera: Tenebrionidae) in the Nebraska Sandhills. Journal of Parasitology 86: 560-565.
  • Clopton, R. E., and R. R. Kula.  1999.  Amoebogregarina nigra n. gen., n. comb. (Apicomplexa: Gregarinidae) From Adult Melanoplus differentialis (Orthoptera: Acrididae) in Southeastern Nebraska. Journal of Parasitology 85: 321-325.
  • Clopton, R. E.  1999.  Revision of the genus Xiphocephalus and description ofXiphocephalus ellisi n. sp.(Apicomplexa: Eugregarinida: Stylocephalidae) fromEleodes opacus (Coleoptera: Tenebrionidae) in the western Nebraska Sandhills.  Journal of Parasitology 85.  84-89.
  • Lucarotti, C. J., T. L. Leclerc, and Clopton, R. E.  1998.  Incidence of Leidyana canadensis (Apicomplexa: Eugregarinida) in Lambdina fiscellaria fiscellaria larvae (Lepidoptoera: Geometridae).  Canadian Entomologist 130: 583-594.
  • Clopton, R. E.  1998.  Revision of the genus Torogregarina and description ofTorogregarina sphinx n. Sp. (Apicomplexa: Eugregarinida) from a Missouri river bank beetle, Bembidion laevigatum (Coleoptera: Carabidae), in southeastern Nebraska. Journal of Parasitology 84: 823-827.
  • Clopton, R. E. and C. J. Lucarotti.  1997.  Leidyana canadensis N. Sp. (Apicomplexa: Eugregarinida) from larval Eastern Hemlock Looper, Lambdina fiscellaria fiscellaria(Lepidoptoera: Geometridae).  Journal of Eukaryotic Microbiology 44: 383-387.
  • Janovy, Jr., S. D. Snyder, and R. E. Clopton.  1997.  Evolutionary constraints on population structure: the parasites of Fundulus zebrinus (Pisces: Cyrinodontidae) in the South Platte River of Nebraska.  Journal of Parasitology 83: 584-592.
  • Clopton, R. E.  1996.  Leidyana migrator n. sp. (Apicomplexa: Leidyanidae) from the Madagacar hissing cockroach, Gromphadorhina portentosa (Insecta: Blattodea).  Invertebrate Biology 114: 271-278.
  • Clopton, R. E. and R. E. Gold.  1996.  Host Specificity of Gregarina blattarum among Five Species of Domiciliary Cockroaches.  Journal of Invertebrate Pathology 67: 219-223.
  • Clopton, R. E.  1995.  Domadracunculus janovyii n. gen., n. sp., (Apicomplexa: Actinocephalidae) described from adults of Ischnura verticalis (Odonata: Zygoptera) in Texas.  Journal of Parasitology 81:256–260.
  • Clopton, R. E. and R. E. Gold.  1995.  Effects of pH on excystation in Gregarina cuneata and Gregarina polymorpha (Eugregarinida: Gregarinidae).  Journal of Eukaryotic Microbiology  42: 540-544.
  • Percival, T. J., R. E. Clopton and J. Janovy, Jr.  1995.  Hoplorhynchus acanthatholius N. Sp. and Steganorhynchus dunwoodii N. Gen. N. Sp. (Apicomplexa: Eugregarinorida) described from Coenagrionid Damselflies.  Journal of Eukaryotic Microbiology  42: 406-410.
  • Janovy, Jr., R. E. Clopton, D. A. Clopton, S. Snyder, A. Efting and L. Krebs.  1994 Species density distributions as null models for ecologically significant interactions of parasite species in an assemblage.  Ecological Modeling 77: 189-196.
  • Clopton, R. E. and R. E. Gold.  1993.  Distribution and Seasonal and diurnal activity patterns of Eutrombicula alfreddugesi (Acari: Trombiculidae) in a forest edge ecosystem. Journal of Medical Entomology 30: 47–53.
  • Clopton, R. E. and J. Janovy, Jr. 1993  Developmental niche structure in the gregarine assemblage parasitizing Tenebrio molitor. Journal of Parasitology 79:701–709.
  • Clopton R. E., T. J. Percival  and John Janovy, Jr.  1993.  Nubenocephalus nebraskensis N. Gen., N. Sp. (Apicomplexa: Actinocephalidae) described from Argia bipunctulata (Odonata: Zygoptera).  Journal of Parasitology 79: 533–537.
  • Ferdig, M. T., M. A. McDowell, R. E. Clopton and Janovy, J. Jr.  1993.  Patterns of morphological variation of Salsuginus yutanensis (Monogenera: Ancyrocephalidae) over space and time.  Journal of Parasitology 79: 744-750.
  • Clopton, R. E. and R. E. Gold.  1992. Bite-Count evaluation of the repellency of N,N-Diethyl-3-methylbenzamide to larval Eutrombicula alfreddugesi (Acari: Trombiculidae).  Journal of Medical Entomology 29: 858-863.
  • Clopton R. E., T. J. Percival and John Janovy, Jr.  1992. Host Stadium Specificity in the gregarine assemblage parasitizing Tenebrio molitor.  Journal of Parasitology 78: 334–337.
  • Clopton R. E., T. J. Percival and John Janovy, Jr.  1992.  Gregarina coronata N. Sp. (Apicomplexa: Eugregarinida) described from adults of the Southern Corn Rootworm, Diabrotica undecimpunctata howardi.  Journal of Protozoology 39: 417–420.
  • Janovy, Jr., R. E. Clopton and T. J. Percival.   1992.  The Roles of ecological and evolutionary influences in providing structure to parasite species assemblages.  Journal of Parasitology 78: 630-640.
  • Clopton R. E., T. J. Percival and John Janovy, Jr.  1991.  Gregarina niphandrodes N. Sp. (Apicomplexa: Eugregarinorida) from adult Tenebrio molitor (L.) with oocyst descriptions of other gregarine parasites of the Yellow Mealworm.  Journal of Protozoology 38:472–483.
  • Clopton R. E.  1991.  A review of the Scaritinid Beetles (Coleoptera: Carabidae) of Nebraska.  Transactions of the Nebraska Academy of Science 18: 53–65.