Philip S. Ward, Entomologist, USA.

  • Research Interests: Systematics and evolutionary biology of ants; genetic structure of social insect populations.
  • Department of Entomology and Nematology, University of California at Davis, Davis, CA 95616, USA.

Taxon names authored

(List may be incomplete)



(List may be incomplete)


  • Ward, P. S., Hebert, P., Harmsen, R. 1973. A note on the distribution of Oeneis chryxus strigulosa (Lepidoptera: Satyridae). Canadian Entomologist, 105: 1373–1374.


  • Harmsen, R., Hebert, P., Ward, P. S. 1974. On the origin of austral elements in the moth fauna of south-eastern Ontario, including a number of species new for Canada. Journal of Research on Lepidoptera, 12: 127–134.
  • Ward, P. S., Harmsen, R., Hebert, P. 1974. An annotated checklist of the macroheterocera of south-eastern Ontario. Journal of Research on Lepidoptera, 13: 23–42.
  • Hebert, P., Ward, P. S., Harmsen, R. 1974. Diffuse competition in Lepidoptera. Nature, 252: 389–391.


  • Hebert, P., Ward, P. S., Harmsen, R. 1975. Competition and species abundance – Reply. Nature, 257: 160–161.


  • Ward, P. S. 1976. Probing ponerine ants. Australian Natural History, l8: 384–387.


  • Ward, P. S. 1980. A systematic revision of the Rhytidoponera impressa group (Hymenoptera: Formicidae) in Australia and New Guinea. Aust. J. Zool. 28: 475–498.
  • Ward, P. S. 1980. Genetic variation and population differentiation in the Rhytidoponera impressa group, a species complex of ponerine ants (Hymenoptera: Formicidae). Evolution, 34: 1060–1076.


  • Ward, P. S., Taylor, R. W. 1981. Allozyme variation, colony structure, and genetic relatedness in the primitive ant Nothomyrmecia macrops Clark (Hymenoptera: Formicidae). Journal of the Australian Entomological Society, 20: 177–183.
  • Ward, P. S. 1981. Ecology and life history of the Rhytidoponera impressa group (Hymenoptera: Formicidae). I. Habitats, nest sites, and foraging behavior. Psyche, 88: 89–108.
  • Ward, P. S. 1981. Ecology and life history of the Rhytidoponera impressa group (Hymenoptera: Formicidae). II. Colony origin, seasonal cycles, and reproduction. Psyche, 88: 109–126.


  • Ward, P. S., Boussy, I. A., Swincer, D. E. 1982. Electrophoretic detection of enzyme polymorphism and differentiation in three species of spider mites (Tetranychus) (Acari: Tetranychidae). Annals of the Entomological Society of America, 75: 595–598.


  • Ward, P. S. 1983. Genetic relatedness and colony organization in a species complex of ponerine ants. I. Genotypic and phenotypic composition of colonies. Behavioral Ecology and Sociobiology, 12: 285–299.
  • Ward, P. S. 1983. Genetic relatedness and colony organization in a species complex of ponerine ants. II. Patterns of sex ratio investment. Behavioral Ecology and Sociobiology, 12: 301–307.




  • Ward, P. S. 1985. The Nearctic species of the genus Pseudomyrmex (Hymenoptera: Formicidae). Quaest. Entomol. 21: 209–246.


  • Ward, P. S. 1988. Mesic elements in the western Nearctic ant fauna: taxonomic and biological notes on Amblyopone, Proceratium, and Smithistruma (Hymenoptera: Formicidae). J. Kans. Entomol. Soc. 61: 102–124


  • Ward, P. S. 1989. Systematic studies on pseudomyrmecine ants: revision of the Pseudomyrmex oculatus and P. subtilissimus species groups, with taxonomic comments on other species. Quaest. Entomol. 25: 393–468


  • Ward, P. S. 1990. The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): generic revision and relationship to other formicids. Systematic Entomology, 15: 449–489.


  • Ward, P. S. 1991. Phylogenetic analysis of pseudomyrmecine ants associated with domatia-bearing plants. pp. 335–352 in: Huxley, C.R., Cutler, D.F. (eds.) Ant-plant interactions. Oxford: Oxford University Press, xviii + 601 pp.


  • Ward, P. S. 1992. Ants of the genus Pseudomyrmex (Hymenoptera: Formicidae) from Dominican amber, with a synopsis of the extant Antillean species. Psyche (Camb.) 99: 55–85
  • Baroni Urbani, C.; Bolton, B.; Ward, P. S. 1992. The internal phylogeny of ants (Hymenoptera: Formicidae). Systematic Entomology, 17: 301–329.


  • Ward, P. S. 1993. Systematic studies on Pseudomyrmex acacia-ants (Hymenoptera: Formicidae: Pseudomyrmecinae). J. Hym. Res. 2: 117–168.




  • Olson, D. M.; Ward, P. S. 1996. The ant fauna (Hymenoptera: Formicidae) of Kirindy Forest (tropical dry forest) in western Madagascar. pp. 161–164 in Ganzhorn, J. U., Sorg, J.-P. (eds.) Ecology and economy of a tropical dry forest in Madagascar. Primate Report 46-1. Göttingen: German Primate Center (DPZ), 382 pp.
  • Ward, P. S. 1996. A new workerless social parasite in the ant genus Pseudomyrmex (Hymenoptera: Formicidae), with a discussion of the origin of social parasitism in ants. Systematic Entomology, 21: 253–263.
  • Ward, P. S.; Bolton, B.; Shattuck, S. O.; Brown, W. L., Jr. 1996. A bibliography of ant systematics. Univ. Calif. Publ. Entomol. 116: 1–417.


  • Ward, P.S. 1999: Deceptive similarity in army ants of the genus Neivamyrmex (Hymenoptera: Formicidae): taxonomy, distribution and biology of N. californicus (Mayr) and N. nigrescens (Cresson). Journal of Hymenoptera research 8(1): 74-97. BHL Reference page
  • Brady, S. G.; Gadau, J.; Ward, P. S. 1999. Is the ant genus Camponotus paraphyletic? [Abstract.] pp. 27–28 in: 4th International Hymenopterists Conference, 6–11 January 1999, Canberra, Australia. Program and abstracts. Glen Osmond, South Australia: 4th International Hymenopterists Conference, 96 pp.
  • Gadau, J.; Brady, S. G.; Ward, P. S. 1999. Systematics, distribution, and ecology of an endemic California Camponotus quercicola (Hymenoptera: Formicidae). Ann. Entomol. Soc. Am. 92(4): 514–522.
  • Ward, P. S. 1999. Systematics, biogeography and host plant associations of the Pseudomyrmex viduus group (Hymenoptera: Formicidae), Triplaris- and Tachigali-inhabiting ants. Zool. J. Linn. Soc. 126: 451–540


  • Ward, P. S. 2000. On the identity of Pheidole vaslitii Pergande (Hymenoptera: Formicidae), a neglected ant from Baja California. J. Hym. Res. 9: 85–98


  • Ward, P. S. 2001. Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera: Formicidae) in the Oriental and Australian regions. Invertebr. Taxon. 15: 589–665.


  • Ward, P., and Brady, S. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics, 17 (3): 361–386.




  • Ward, P.S. 2006. The ant genus Tetraponera in the Afrotropical region: synopsis of species groups and revision of the T. ambigua-group. Myrmecologische Nachrichten, 8: 119–130.


  • Ward, P.S. 2007. Phylogeny, classification, and species-level taxonomy of ants. Zootaxa, 1668: 549–563.
  • Ward, P.S. 2007. The ant genus Leptanilloides: discovery of the male and evaluation of phylogenetic relationships based on DNA sequence data (pp. 637–649). In Snelling, R.R., Fisher, B.L. & Ward, P.S. (eds). Advances in ant systematics: homage to E.O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute 80: 690 pp.


  • Ward, P.S. 2009. The ant genus Tetraponera in the Afrotropical region: the T. grandidieri group. Journal of Hymenoptera Research, 18: 285–304.
  • Ward, P.S. & Brady, S.G. 2009. Rediscovery of the ant genus Amyrmex Kusnezov and its transfer from Dolichoderinae to Leptanilloidinae. Zootaxa, 2063: 46–54.


  • Ward, P.S., Brady, S.G., Fisher, B.L. & Schultz, T.R. 2010. Phylogeny and biogeography of Dolichoderinae ants: effects of data partitioning and relict taxa on historical inference. Systematic Biology, 59: 342–362.
  • Lucky, A., Ward, P. S. 2010. Taxonomic revision of the ant genus Leptomyrmex Mayr (Hymenoptera: Formicidae). Zootaxa, 2688: 1–67. [Pdf file]


  • Ward, P. S. 2011. Integrating molecular phylogenetic results into ant taxonomy (Hymenoptera: Formicidae). Myrmecological News, 15: 21–29.
  • Roura-Pascual, N., Hui, C., Ikeda, T., Leday, G., Richardson, D. M., Carpintero, S., Espadaler, X., Gómez, C., Guénard, B., Hartley, S., Krushelnycky, P., Lester, P. J., McGeoch, M. A., Menke, S. B., Pedersen, J. S., Pitt, J. P. W., Reyes, J., Sanders, N. J., Suarez, A. V., Touyama, Y., Ward, D., Ward, P. S., Worner, S. P. 2011. The relative roles of climatic suitability and anthropogenic influence in determining the pattern of spread in a global invader. Proceedings of the National Academy of Sciences U. S. A. 108: 220–225.


  • Ward, P. S., Sumnicht, T. P. 2012. Molecular and morphological evidence for three sympatric species of Leptanilla (Hymenoptera: Formicidae) on the Greek island of Rhodes. Myrmecological News.


  • Ward, P.S., Brady, S.G., Fisher, B.L. & Schultz, T.R. 2014 [2015]. The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40(1): 61–81. DOI: 10.1111/syen.12090 Reference page
  • Brady, S.G., Fisher, B.L., Schultz, T.R. & Ward, P.S. 2014. The rise of army ants and their relatives: diversification of specialized predatory doryline ants. BMC Evolutionary Biology 14: 93.
  • Ward, P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 23–43.


  • Blaimer, B. B., Brady, S. G., Schultz, T. R., Lloyd, M. W., Fisher, B. L., & Ward, P. S. 2015. Phylogenomic methods outperform traditional multi-locus approaches in resolving deep evolutionary history: a case study of formicine ants. BMC evolutionary biology 15: 271.
  • Chomicki, G., Ward, P. S., Renner, S. S. 2015. Macroevolutionary assembly of ant/plant symbioses: Pseudomyrmex ants and their ant-housing plants in the Neotropics. In Proceedings of the Royal Society B 282: 20152200.




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