Philip S. Ward

Philip S. Ward


Office: 381A Briggs Hall
Lab: 381B Briggs Hall
Phone: (530) 752-0486
Fax: (530) 752-1537
B.Sc. Biology, Queens University, Canada (1973)
Ph.D. Zoology, University of Sydney, Australia (1979)
25% Instruction and Research
75% Organized Research
Systematics, biogeography and evolution of ants; ant-plant mutualisms; phylogeny and speciation.
Research Interests:
Systematics and evolutionary biology of ants; genetic structure of social insect populations.


Baroni Urbani, C.; Bolton, B.; Ward, P. S. 1992.  The internal phylogeny of ants (Hymenoptera: Formicidae). Syst. Entomol. 17: 301-329

Brady, S. G.; Gadau, J.; Ward, P. S. 1999.  Is the ant genus Camponotus paraphyletic? [Abstract.] Pp. 27-28 in: 4th International Hymenopterists Conference, 6-11th 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

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. 1980a.  A systematic revision of the Rhytidoponera impressa group (Hymenoptera: Formicidae) in Australia and New Guinea. Aust. J. Zool. 28: 475-498

Ward, P. S. 1984.  A revision of the ant genus Rhytidoponera (Hymenoptera: Formicidae) in New Caledonia. Aust. J. Zool. 32: 131-175

Ward, P. S. 1985b.  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. 1989a.  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. Syst. Entomol. 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

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

Ward, P. S. 1994.  Adetomyrma, an enigmatic new ant genus from Madagascar (Hymenoptera: Formicidae), and its implications for ant phylogeny. Syst. Entomol. 19: 159-175

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. Syst. Entomol. 21: 253-263

Ward, P. S. 1999a.  Deceptive similarity in army ants of the genus Neivamyrmex (Hymenoptera: Formicidae): taxonomy, distribution and biology of N. californicus (Mayr) and N. nigrescens (Cresson).  J. Hym. Res. 8: 74-97

Ward, P. S. 1999b.  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. 2000a.  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.S. 2005. A synoptic review of the ants of California. Zootaxa 936: 1-68.

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. 2007a. Phylogeny, classification, and species-level taxonomy of ants. Zootaxa 1668: 549-563.

Ward, P.S. 2007b. 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.; Bolton, B.; Shattuck, S. O.; Brown, W. L., Jr. 1996.  A bibliography of ant systematics. Univ. Calif. Publ. Entomol. 116: 1-417

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. & 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.

Ward, P.S. & Downie, D.A. 2005. The ant subfamily Pseudomyrmecinae: phylogeny and evolution of big-eyed arboreal ants. Systematic Entomology 30: 310-335.

Ward, P.S. & Taylor, R.W. 1981. Allozyme variation, colony structure and genetic relatedness in the primitive ant Nothomyrmecia macrops Clark. Journal of the Australian Entomological Society 20: 177-183. [24.viii.1981.]

Bolton B, Alpert G, Ward PS, Naskrecki P. 2007. [CD-ROM] Bolton's Catalogue of the Ants of the World. Harvard University Press.