Department of Entomology


John Heratyucr entomology

Professor of Entomology

Location: Entomology 138
(951) 827-6351


Research Specialization - Our research focuses on the systematics, phylogeny and biogeography of the Chalcidoidea (Hymenoptera). Chalcidoid wasps rank numerically among the largest groups of insects, with estimates of as many as 100,000 species; however, the fauna is poorly known. Most are specialized parasites, and the majority of successful biological control projects have utilized these minute wasps to achieve partial or complete control of insect pests. One area of our research is on the Eucharitidae, a specialized group of ant parasities. The taxonomy of this group is poorly understood, and presently, only a small proportion of the known species can be identified. The larvae of Eucharitidae exhibit several peculiar behaviors associated with gaining access to the ant host, and studies on higher classification using cladistic methodology have led to a greater understanding of the evolution of behavioral patterns within the family, This knowledge has proved useful in studies on other groups of Hymenoptera that parasitize eusocial insects and for postulating a biogeographic hypothesis for the family. A second area of research emphasizes the systematics of the Aphelinidae, which are generally parasites of aphids, whiteflies and scale insects. Most analyses of relationships have been based on internal and external morphological traits, but molecular techniques are now being applied to understanding the higher phylogeny of the Chalcidoidea and the relationships among species of the genus Encarsia (Aphelinidae). Other research interests include studies of the diversity of Hymenoptera on the Galapagos Islands, internal studies of the skeleto-musculature of Hymenoptera and parasitoid interactions with leafmining moths of the family Gracillariidae. All of our studies incorporate morphological, biological or molecular information into analyses that are used to formulate hypotheses of phylogenetic relationships and the evolution of behavioral patterns. The evolution of host associations, an area of paramount importance to biological control programs, is central to all of our studies. This is a fascinating area of research demonstrating the utility and impact of systematics to almost every area of science.


BSc Biology 1976
University of Guelph, Ontario, Canada
MSc Entomology/Systematics 1985
University of Guelph, Ontario, Canada
PhD Entomology/Systematics 1990
Texas A&M University


1993-1994 - Smithsonian Institute Postdoctoral Fellowship
1991 - Honorary Research Associate, Royal Ontario Museum,

Research Area

Morphological and molecular systematics of Chalcidoidea Cladistic methodology; Biological control Evolutionary biology.


Heraty, J.M., Burks, R.A., Mbanyana, N., van Noort, S. 2018. Morphology and life history of an ant parasitoid, Psilocharis afra (Hymenoptera: Eucharitidae). Zootaxa 4482: 491-510.

Burks, R.A., Krogmann, L., J.M. Heraty. 2018. Simultaneous discovery and taxonomic placement of new extant and fossil genera of Herbertiinae (Hymenoptera: Chlacidoidea: Pteromalidae). Insect Systematics and Diversity 2: 1-7 doi: 10.1093/isd/ixy012

Chien, I., J.M. Heraty. 2017 Come and gone: Description of a new species of Eucharitidae (Hymenoptera) attacking Solenopsis (Hymenoptera: Formicidae) in Texas. Insect Systematics and Diversity 2: 1-7. doi: 10.1093/isd/izz012

Burks, R.A., Heraty, J.M., Dominguez, C.J. Mottern. 2018. Complex dviersity in a mainly tropical group of ant parasitoids: Revision of the Orasema stramineipes species group (Hymenoptera: Chalcidoidea: Eucharitidae). Zootaxa 44:107 pp.

Janšta, P., Cruaud, A., Delvare, G., Genson, G., Heraty, J., Krízková, B., Rasplus, R.-Y. 2018. Torymidae (Hymenoptera, Chalcidoidea) revised: molecular phylogeny, circumscription and reclassification of the family with discussion of its biogeography and evolution of life-history traits. Cladistics DOI: 10.1111/cla. 12228

Peters, R.S., Niehuis, O., Gunkel, S., Bläser, M., Mayer, C., Podsiadlowski, L., Kozlov, A., Donath, A., van Noort, S. Liu, S., Zhou, X., Misof, M., Heraty, J., Krogmann, L. 2018. Transcriptome sequence-based phylogeny of chalcidoid wasps (Hymenoptera: Chalcidoidea) reveals a history of rapid radiations, convergence, and evolutionary success. Molecular Phylogenetics and Evolution 120: 286-296.

Hash, J.M., Brown, B.V. and Heraty, J.M. 2018. Host association and biogeographic patterns in the diverse millipede-parasitoid genus Myriophora Brown (Diptera: Phoridae). Cladistics 34: 93-112.

Burks, R.A., Mottern, J.L., Dominguez, C., Heacox, S. and J.M. Heraty. 2017. Biting the bullet: Revisionary notes on the Oraseminae of the Old World (HymenopteraL Chalcidoidea: Eucharitidae). Journal of Hymenoptera Research 55: 139-188.

Peters, R.S., Korgmann, L., Mayer, C., Donath, A., Gunkel, S., Muesemanm, K., Kozlov, A., Podsiadlowski, L., Petersen, M., Lanfear. R., Diez, P., Heraty, J.M., Kjer, K, K., Klopfstein, S., Meier, R., Polidori, C., Schmitt, T., Liu, S., Zhou, X., Wappler, T., Rust, J., Misof, B. and O. Nieuis 2017. Evolutionary history of the sawflies, wasps, ants, and bees. Current Biology 27, 1013-1018.

Hopper, K.R., Lanier, K., Rhoades, J.H., Coutinot, D., Mercadier, G., Ramaulde, N., Roche, M., Woolley, J.B., and J.M. Heraty. 2017. Host specifically of Aphelinus species considered for introduction to control Diuraphis noxia. Biological Control 107: 21-32.

Hash, J.M., Millar, J.G., Heraty, J.M., Harwood, J.F. and Brown, B.V. 2017. Milipede defensive compounds are a double-edged sword: Natural history of the millipede parasitic genus Myriophora (Diptera: Phoridae). Journal of Chemcial Ecology doi: 10.1007/s10886-016-0815-7

Herreid, J.S. Heraty, J.M. 2017. Hitchhikers at the dinner table: exploitation of extrafloral nectaries by a monophyletic group of ant parasitoids (Hymenoptera: Eucharitidae). Systematic Entomology 42: 204-229.

Morse, J.G., Rugman-Jones, P., Woolley, J.B., Heraty, J.M. Triapitsyn, S.V., Robinson, L.J., Hofshi, R. and Stouthamer, R. 2016. Armored scales and their parasitoids on commercial avocados in California and imported fruit from Mexico. Journal of Economic Entomology 109: 2032-2042. doi.org/10.1093/jee/tow155

Murray, E. and Heraty, J.M. 2016. Invading Africa: a novel transoceanic dispersal by a New World ant parasitoid. Journal of Biogeography 43, 1750-1761.

Torréns, J., Heraty, J.M., Murray, E. and Fidalgo, P. 2016. Biology and phylogenetic placement of a new species of Lasiokapala Ashmead from Argentina (Hymenoptera: Eucharitidae). Systematic Entomology 41, 596-606.

Feng, L., Bhanu, B. and Heraty, J.M. 2016. A software system for automated identification and retrieval of moth images based on wing attributes. Pattern Recognition 51: 225-241.

Heraty, J.M., Mottern, J. and Peeters, C. 2015. A new species of Schizaspidia, with discussion of the phylogenetic utility of immature stages for assessing relationships among eucharitid parasitoids of ants. Annals of the Entomological Society of America DOI: 10.1093/aesa/sav062

Burks, R.A., Mottern, J. and Heraty, J.M. 2015. Revision of the Orasem festiva species group (HymenopteraL Chalcidoidea: Eucharitidae). Zootaxa 3972 (4): 521-534.

Burks, R.A., Heraty, J.M., Pinto, J.D. and Grimaldi, D. 2015. Small but not ephemeral: newly discovered species of Aphelinidae and Trichogrammatidae (Insecta: Hymenoptera: Chalcidoidea) from Eocene amber/ Systematic Entomology. DOI: 10.111/syen. 12124

Burks, R.A. and Heraty, J.M. 2015. Subforaminal bridges in Hymenoptera (Insecta), with a focus on Chalcidoidea. Arthropod Structure and Development 44: 173-194.

Mottern, J.L. and Heraty, J.M. 2014. Revision of a cryptic species complex in the genus Cales (Hymenoptera, Chalcidoidea, Aphelinidae). Systematic Entomology 39: 354-379.

Mottern, J.L. and Heraty, J.M. 2014. The dead can talk: Museum specimens show the origins of a cryptic species used in biological control. Biological Control 71: 30-39.

Heraty, J.M. and Murray, E. 2013. The life history of Pseudometagea schwarzii, with a discussion of the evolution of endoparasitism and koinobiosis in a specialized group of chalcid wasps. Journal of Hymenoptera Research 35: 1-15.

Heraty, J.M. Burks, R.A., Cruaud, A., Gibson, G.A.P., et al. (27 authors). 2013. A Phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics, 29: 466-542. doi/10.1111/cla.12006

Klopfstein, S., Vilhelmsen, L. Heraty, J., Sharkey, M. and Ronquist, F. 2013. The hymenopteran tree of life: evidence from protein-coding genes and objectively aligned ribosomal data. PLoS one 8: e69344. 23 pp.

Murray, E., Carmichael, A.E. and Heraty, J.M. 2013. Ancient host shifts followed by host conservatism in a group of ant parasitoids. Proceedings of the Royals Society, B. 280: 0130495. (9 pages)

Torréns, J. and Heraty, J.M. 2013. A new genus of Eucharitidae (Hymenoptera: Chalcidoidea), with notes on life history and immature stages. Zootaxa 3630: 347-358.

Torréns, J. and Heraty, J.M. 2012. Description of the species of Dicoelothorax Ashmead (Chalcidoidea, Eucharitidae) and biology of D. platycerus Ashmead. Zookeys 165: 33-46.

Kondo, T., Quintero Q, E. M., Campuzano, M., Wyckhuys, K. A. G. and Heraty, J.M. 2012. First report of Tamarixia radiata (Waterston) (Hymenoptera: Eulophidae), a paraditoid of the Asian citrus psyllid Diaphorina citri Kuwayama (Hemiptera Psyllidae), in the Department of Valle del Cauca, Colombia. Boletín del Museo de Entomología de la Universidad de Valle 13: 48-51.

Rugman-Jones, P.F., Hoddle, M.S., Amrich, R., Heraty, J.M., Stouthamer-Ingel and C.E., Stouthamer, R.S. 2012. Phylogeographic structure, outbreeding depression, and reluctant virgin oviposition in the bean thrips, Caliothrips fasciatus (Pergande) (Thysanoptera: Thripidae), in California. Bulletin of Entomological Research 102: 698-709.

Kim, J. and Heraty, J. 2012. A phylogenetic analysis of the genera of Aphelininae (Hymenoptera: Aphelinidae), with a generic key and descriptions of a new taxa. Systematic Entomology 37, 497-549.

Carey, B., Visscher, K. and Heraty, J. 2012. Extrafloral nectaries for gaining access to an ant host by the parasitoid Orasema simulatrix (Hymenoptera: Eucharitidae). Journal of Hymenoptera Research 27: 47-65.

Munro, J.B., Heraty, J., Burks, R.A., Hawks, D., Mottern, J.L., Cruaud, A., Rasplus., J.-Y. and Jansta, P. 2011. A molecular phylogeny of the Chalcidoidea (Hymenoptera). PLoS One 6: e27023.

Sharkey, M.J., Carpenter, J.C., Vilhelmsen, L., Heraty, J.M., Liljeblad, J., Dowling, A.P.G., Schulmeister, S., Murray, D., Deans, A., Ronquist, F., Korgmann, L., Wheeler, W.C. 2011. Phylogenetic relationships among superfamilies of Hymenoptera. Cladistics 27: 1-33.

Heraty, J.M., Ronquist, F., Carpenter, J.C., Hawks, D., Schulmeister, S., Dowling, A., Murray, D., Munro, J.B., Schiff, N., Wheeler, W.C., and Sharkey, M., 2011. Hymenopteran relationshipsL Structure of a megaradiation. Molecular Phylogenetics and Evolution 60: 73-88.

Mottern, J., Heraty, J.M. and Hartop, E. 2011. Calesinae (Hymenoptera: Chalcidoidea): morphology of an enigmatic taxon with a review of species. Systematic Entomology 36: 267-28.

Burks, R.A., Heraty, J.M., Gebiola, M. and Hansson, C. 2011. Combined molecular and morphological phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with focus on the subfamily Entedoninae. Cladistics 27: 1-25.

Varone, L. Heraty, J.M., Calcaterra, L.A. 2010. Distribution, abundance and persistence of species of Orasema (Hym: Eucharitidae) parasitic on fire ants in South America. Biological Control 55: 72-78.

Romero, C. and J.M. Heraty. 2010. Ultrastructure of scutellar sensilla in Aphytis melinus (Hymenoptera: Aphelinidae) and morphological variation across Chalcidoidea. Journal of Hymenoptera Research 19: 164-183.

Heraty, J.M. and D.C. Darling. 2009. Fossils Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea) from Baltic Amber. Zootaxa 2306: 1-16.

Heraty, J.M. Heraty and J. Torréns. 2009. A new species of Pseudochalcura (Hymenoptera: Eucharitidae), with a review of antennal morphology from a phylogenetic perspective. Zookeys. 20: 215-231.

Bhanu, B, R. Li, J. Heraty and E. Murray. 2009. Automated classification of skippers based on parts representation. American Entomologist 54: 228-231.

Kirkland, C.A., R.A. Burks, D.H. Headrick and J.M. Heraty. 2009. A review of Hadrotrichodes waukheon LaSalle (Hymenoptera: Eulophidae): A possible biological control agent of Marmara gulosa Guillén & Davis (Lepidoptera: Gracillariidae) in San Joaquin Valley Citrus. Proceedings of the Entomological Society of Washington 111: 7-11.

Hoddle, M.S., J.M. Heraty, P.F. Rugman-Jones, L.A. Mound and R.S. Stouthamer. 2008. Relationships among species of Scirtothrips Shull 1909 (Thysanoptera: Thripidae, Thripinae) using molecular and morphological data. Annals of the Entomological Society of America 101: 491-500.

Wang, Q., J. Millar, J. Mottern, J. Heraty, S. Triapitsyn and T.D. Paine. 2008. Selection and collection of a strain of the egg parasitoid Avetianella longoi that attacks Phoracantha recurva in Australia. Journal of Economic Entomology 101: 1771-1778.

Torréns, J., Heraty, J.M., Fidalgo, P. 2008. Biology and description of a new species of Lophyrocera Cameron (Hymenoptera: Eucharitidae from Argentina. Zootaxa 1871: 56-62.

Buffington, M., F. Nylander and J.M. Heraty. 2007. The phylogeny, evolution of Figitidae (Hymenoptera: Cynipoidea). Cladistics 23: 1-19.

Guillén, M., J. M. Heraty and A. Urbaneja. 2007. Fecundity and development of Cirrospilus coachellae (Hymenoptera: Eulophidae), a parasitoid of Marmara gulosa (Lepidoptera: Gracillariidae). Journal of Economic Entomology 100: 664-669.

Guillén, M., R.F. Luck and J.M. Heraty. 2007. Preferred host stages, clutch size and sex allocation by Cirrospilus coachellae (Hymenoptera: Eulophidae), a parasitoid of Marmara gulosa (Lepidoptera: Gracillariidae).
Biocontrol. 40: 264-272.

Hansen, I.A., D.H. Sieglaff, J.B. Munro, S.-H. Shiao, J. Cruz, I.W. Lee, J. Heraty and A.S. Raikhel. 2007. Forkhead transcription factors regulate mosquito reproduction. Insect Biochemistry and Molecular Biology 37: 985-997.

Heraty, J.M. and D.C. Darling. 2007. A new genus and species of Perilampidae (Hymenoptera: Chalcidoidea) with uncertain placement within the family. Proceedings of the Entomological Society of Ontario 138: 33-47.

Heraty, J.M., J.B. Woolley, K.M. Hopper, D.L. Hawks, J.-W. Kim and M. Buffington. 2007. Phylogenetic relationships of cryptic species in the Aphelinus varipes complex (Hymenoptera: Aphelinidae). Molecular Phylogenetics and Evolution. 45: 480-493.

Owen, A., J. George, J. Pinto and J.M. Heraty. 2007. A molecular phylogeny of the Trichogrammatidae (Hymenoptera: Chalcidoidea), with an evaluation of the utility of their male genitalia for higher level classification. Systematic Entomology 32: 227-251.

Torréns, J., J.M. Heraty and P. Fidalgo. 2007. Biology and description of a new species of Laurella Heraty (Hymenoptera: Eucharitidae) from Argentina. Proceedings of the Entomological Society of Washington 109: 45-51.

Van Dam, A.R., M.H. Van Dam and J.M. Heraty. 2007. A comparison of pyrethrum fogging and sweep netting of Hymenoptera in southern California chaparral. Journal of Hymenoptera Research 16: 192-205.

Perez-Lachaud, G., J.M. Heraty, A. Carmichael, and J.P. Lachaud. 2006. Biology and behavior of Kapala (Hymenoptera: Eucharitidae) attacking Ectatomma and Gnamptogenys (Formicidae: Ectatomminae) in Chiapas, Mexico. Annals of the Entomological Society of America 99: 567-576.

Smith, A.B.T., D.C. Hawks and J.M. Heraty. 2006. An overview of the classification and evolution of the major scarab beetle clades (Coleoptera: Scarabaeoidea) based on preliminary molecular analyses. Coleopterists Society Monograph 5: 35-46.

Triapitsyn, S.V., D.B. Vickerman, J.M. Heraty and G.A. Logarzo. 2006. A new species of Gonatocerus (Hymenoptera: Mymaridae) parasitic on proconine sharpshooters (Hemiptera: Cicadellidae) in the New World. Zootaxa 1158: 55-67.

Gillespie, J.J., J.B. Munro, J.M. Heraty, M.J. Yoder, A.K. Owen and A. Carmichael. 2005. A secondary structural model of the 28S rRNA expansion segments D2 and D3 from chalcidoid wasps (Hymenoptera: Chalcidoidea): Implications for multiple sequence alignment and phylogeny reconstruction. Molecular Biology and Evolution 22: 1593-1608.

Heraty, J. M. 2005. A revision of the genus Isomerala (Hymenoptera: Eucharitidae). Acta Societatis Zoologicae Bohemicae 69: 137-145.
Guillén, M. and J.M. Heraty. 2004. Instar differences in Marmara gulosa Guillén and Davis (Lepidoptera: Gracillariidae). Annals of the Entomological Society of America 97: 1227-1232.

Heraty, J., D. Hawks, J.S. Kostecki and A. Carmichael. 2004. Phylogeny and behaviour of the Gollumiellinae, a new subfamily of the ant-parasitic Eucharitidae (Hymenoptera: Chalcidoidea). Systematic Entomology 29: 544-559.

Guillén, M., J.M. Heraty and R.F. Luck. 2003. Seasonal variation and infestation incidence by Marmara gulosa Guillén and Davis (Lepidoptera: Gracillariidae) on grapefruit in the Coachella Valley of California (USA). Journal of Economic Entomology 96: 577-583.

Heraty, J.M. and D.J. Quicke. 2003. Phylogenetic implications of ovipositor structure in Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea). Journal of Natural History 37: 1751-1764.

Heraty, J.M. 2003. Three new species of Gollumiella Hedqvist (Hymenoptera: Eucharitidae). Zootaxa 497: 1-10.

Burks, R.D. and J.M. Heraty. 2002.
Morphometric analysis of four species of Trichogramma (Hymenoptera, Trichogrammatidae) attacking codling moth and other tortricid pests in North America. Journal of Hymenoptera Research 11: 167-187.

Gates, M.W., J.M. Heraty, M.E. Schauff, D.L. Wagner, J.B. Whitfield and D.B. Wahl. 2002. Survey of the parasitic Hymenoptera on leafminers in California. Journal of Hymenoptera Research 11: 213-270.

Babcock, C.S., J.M. Heraty, P.J. De Barro, F. Driver, and S. Schmidt. 2001. Preliminary phylogeny of Encarsia Förster (Hymenoptera: Aphelinidae) based on morphology and 28S rDNA. Molecular Phylogenetics and Evolution 18: 306-323.

Babcock, C.S. and J.M. Heraty. 2000. Molecular markers for the discrimination of Encarsia formosa and Encarsia luteola (Hymenoptera: Aphelinidae). Annals of the Entomological Society of America 93: 738-744.

Guillén, M., D. Davis, and J.M. Heraty. 2001. A new species of Marmara (Lepidoptera: Gracillariidae) infesting Citrus in the southwestern United States. Proceedings of the Entomological Society of Washington 103: 636-654.

Heraty, J. M. 2000. Phylogenetic relationships of Oraseminae (Hymenoptera: Eucharitidae). Annals of the Entomological Society of America 93: 374-390.

Heraty, J.M. and A.P. Polaszek. 2000. Morphometric analysis and descriptions of selected species in the Encarsia strenua species group (Hymenoptera: Aphelinidae). Journal of Hymenoptera Research 9: 142-169.

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