Philip S. Ward



Philip Ward
Dept. of Entomology and Nematology
University of California, Davis
One Shields Avenue
Davis, CA 95616

Email: psward [at]

My research is motivated by a fascination with insect natural history and the processes that have generated such a diverse array of forms. In the Ward lab our specialty is the systematics and evolutionary biology of ants (Hymenoptera: Formicidae). These creatures, while small in size, are the world’s most prominent and species-rich group of social insects. Numbering more than 14,000 described species—and perhaps as many undescribed taxa—ants have come to occupy almost all major terrestrial habitats, and they often assume keystone roles as predators, scavengers and herbivores. Our objectives are to document and analyze the diversity of ants and to provide insight into the factors responsible for their evolutionary success.

As an ant systematist I engage in two different but complementary activities: (1) species-level taxonomy, and (2) phylogenetics. The former is a neglected but vitally important area of systematics—without the ability to distinguish species many other enterprises in biology are seriously compromised. Much of my own work on species delimitation has focused on the ant subfamily Pseudomyrmecinae. I am also interested in the natural history and distribution of California ant species, and I have been involved in a collaborative project on the systematics and biogeography of Mesoamerican ants. My phylogenetic studies, based primarily on molecular data (initially Sanger sequencing, but now targeted enrichment of UCE loci), have been directed at both the deep history of ants—unraveling the major lineages of the ant tree of life—and at relationships among closely related species.

Research Interests

The Ant Subfamily Pseudomyrmecinae
ANTBIB: A Bibliography of Ant Systematics Literature


  1. Ward, P. S., Hebert, P., Harmsen, R. 1973. A note on the distribution of Oeneis chryxus strigulosa (Lepidoptera: Satyridae). Canadian Entomologist 105: 1373-1374.
  2. 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.
  3. 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.
  4. Hebert, P., Ward, P. S., Harmsen, R. 1974. Diffuse competition in Lepidoptera. Nature 252: 389-391.
  5. Hebert, P., Ward, P. S., Harmsen, R. 1975. Competition and species abundance – Reply. Nature 257: 160-161.
  6. Ward, P. S. 1976. Probing ponerine ants. Australian Natural History l8: 384-387.
  7. Ward, P. S. 1980. A systematic revision of the Rhytidoponera impressa group (Hymenoptera: Formicidae) in Australia and New Guinea. Australian Journal of Zoology 28 :475-498. [PDF file]
  8. 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. [PDF file]
  9. 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.
  10. 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. [PDF file]
  11. 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. [PDF file]
  12. 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.
  13. 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.
  14. 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.
  15. Ward, P. S. 1984. A revision of the ant genus Rhytidoponera (Hymenoptera: Formicidae) in New Caledonia. Australian Journal of Zoology 32: 131-175. [PDF file]
  16. Ward, P. S. 1985. The Nearctic species of the genus Pseudomyrmex (Hymenoptera: Formicidae). Quaestiones Entomologicae 21: 209-246. [PDF file]
  17. Ward, P. S. 1985. Taxonomic congruence and disparity in an insular ant fauna: Rhytidoponera in New Caledonia. Systematic Zoology 34: 140-151. [PDF file]
  18. Tibayrenc, M., P. Ward, A. Moya and F. Ayala. 1986. Natural populations of Trypanosoma cruzi, the agent of Chagas’ disease, have a complex multiclonal structure. Proceedings of the National Academy of Sciences U. S. A. 83: 115-119.
  19. Ward, P. S. 1986. Functional queens in the Australian greenhead ant, Rhytidoponera metallica (Hymenoptera: Formicidae). Psyche 93: 1-12. [PDF file]
  20. Ward, P. S. 1987. Distribution of the introduced Argentine ant (Iridomyrmex humilis) in natural habitats of the lower Sacramento Valley, and its effects on the indigenous ant fauna. Hilgardia 55(2): 1-16. [PDF file]
  21. Ward, P. S. 1988. Mesic elements in the western Nearctic ant fauna: taxonomic and biological notes on AmblyoponeProceratium, and Smithistruma (Hymenoptera: Formicidae). Journal of the Kansas Entomological Society 61: 102-124. [PDF file]
  22. Ward, P. S. 1989. Genetic and social changes associated with ant speciation. In Breed, M. D. & R. E. Page (Eds.). The genetics of social evolution. Boulder, Colorado: Westview Press, pp. 123-148.
  23. 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. Quaestiones Entomologicae 25: 393-468. [PDF file]
  24. Ward, P.S. 1990. The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): generic revision and relationship to other formicids. Systematic Entomology 15: 449-489. [PDF file]
  25. Ward, P.S. 1991. Phylogenetic analysis of pseudomyrmecine ants associated with domatia-bearing plants. In Cutler, D.F. & C.R. Huxley (Eds.) Ant/plant interactions. Oxford: Oxford Univ. Press, pp. 335-352.
  26. Frumhoff, P.C., Ward, P.S. 1992. Individual-level selection, colony-level selection, and the association between polygyny and worker monomorphism in ants. American Naturalist 139: 559-590.
  27. Baroni Urbani, C., Bolton, B., Ward, P.S. 1992. The internal phylogeny of ants (Hymenoptera: Formicidae). Systematic Entomology 17: 301-329.
  28. Ward, P.S. 1992. Ants of the genus Pseudomyrmex (Hymenoptera: Formicidae) from Dominican amber, with a synopsis of the extant Antillean species. Psyche 99: 55-85. [PDF file]
  29. Gullan, P. J., Buckley, R. C., Ward, P. S. 1993. Ant-tended scale insects (Hemiptera: Coccidae: Myzolecanium) within lowland rainforest trees in Papua New Guinea. Journal of Tropical Ecology 9: 81-91. [PDF file]
  30. Ward, P. S. 1993. Systematic studies on Pseudomyrmex acacia-ants (Hymenoptera: Formicidae: Pseudomyrmecinae). Journal of Hymenoptera Research 2: 117-168. [Pdf file]
  31. Ward, P. S. 1994. Adetomyrma, an enigmatic new ant genus from Madagascar (Hymenoptera: Formicidae), and its implications for ant phylogeny. Systematic Entomology 19: 159-175.
  32. Rejmánek, M., Ward, P. S., Webster, G. L., Randall, J. M. 1994. Systematics and biodiversity. [Correspondence.] Trends in Ecology and Evolution 9: 228-229.
  33. 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.
  34. Ward, P. S., Bolton, B. B., Shattuck, S. O., Brown, W. L., Jr. 1996. A bibliography of ant systematics. University of California Publications in Entomology 116: 1-417.
  35. 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. [PDF file]
  36. Ward, P. S. 1997. Ant soldiers are not modified queens. Nature 385: 494-495.
  37. 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). Journal of Hymenoptera Research 8: 74-97.
  38. Gadau, J., Brady, S. G., Ward, P. S. 1999. Systematics, distribution, and ecology of an endemic California Camponotus quercicola (Hymenoptera: Formicidae). Annals of the Entomological Society of America 92: 514-522.
  39. Ward, P. S. 1999b. Systematics, biogeography and host plant associations of the Pseudomyrmex viduus group (Hymenoptera: Formicidae), Triplaris– and Tachigali-inhabiting ants. Zoological Journal of the Linnean Society 126: 451-540. [PDF file]
  40. Wetterer, J. K., Ward, P. S., Wetterer, A. L., Longino, J. T., Trager, J. C., Miller, S. E. 2000. Ants (Hymenoptera: Formicidae) of Santa Cruz Island, California. Bulletin. Southern California Academy of Sciences 99: 25-31.
  41. Ward, P. S. 2000. On the identity of Pheidole vaslitii Pergande (Hymenoptera: Formicidae), a neglected ant from Baja California. Journal of Hymenoptera Research 9: 85-98. [PDF file]
  42. Ward, P. S. 2000. Broad-scale patterns of diversity in leaf litter ant communities. Pp. 99-121 in: Agosti, D., Majer, J. D., Alonso, L. E., Schultz, T. R. (eds.) Ants. Standard methods for measuring and monitoring biodiversity. Washington: Smithsonian Institution Press, xix + 280 pp.
  43. Brady, S. G., Gadau, J., Ward, P. S. 2000. Systematics of the ant genus Camponotus (Hymenoptera: Formicidae): a preliminary analysis using data from the mitochondrial gene cytochrome oxidase I. Pp. 131-139 in: Austin, A. D., Dowton, M. (eds.) Hymenoptera. Evolution, biodiversity and biological control. Collingwood, Victoria: CSIRO Publishing, xi + 468 pp.
  44. Ward, P. S. 2001. Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera: Formicidae) in the Oriental and Australian regions. Invertebrate Taxonomy 15: 589-665. [PDF file]
  45. Gronlund, C. J., Deangelis, M. D., Pruett-Jones, S., Ward, P. S., Coyne, J. A. 2002. Mate grasping in Drosophila pegasa.  Behaviour 139: 545-572.
  46. Johnson, R. A., Ward, P. S. 2002.  Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: a first overview.  Journal of Biogeography 29:1009-1026. [PDF file]
  47. Boulton, A. M., Ward, P. S. 2002.  Ants.  Pp. 112-128 in Case, T. J., Cody, M. L. & Ezcurra, E. (eds.) A new island biogeography of the Sea of Cortés. Oxford: Oxford University Press, xvii + 669 pp
  48. Ward, P. S., Boulton, A. M. 2002.  Checklist of the ants of the Gulf of California islands.  Pp. 545-553 in Case, T. J., Cody, M. L. & Ezcurra, E. (eds.) A new island biogeography of the Sea of Cortés. Oxford: Oxford University Press, xvii + 669 pp.
  49. Ward, P. S., Brady, S. G.  2003.  Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae).  Invertebrate Systematics 17: 361-386. [PDF file]
  50. Ward, P. S.  2003.  Subfamilia Pseudomyrmecinae.  Pp. 331-333 in Fernández, F. (ed.) Introducción a las hormigas de la región Neotropical. Bogotá, Colombia: Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, xxvi + 398 pp. [PDF file]
  51. Kaspari, M., Ward, P. S., Yuan, M. 2004. Energy gradients and the geographic distribution of local ant diversity. Oecologia 140: 407-413. [PDF file]
  52. Ward, P. S., Downie, D. A.  2005.  The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants. Systematic Entomology 30: 310-335. [PDF file]
  53. Ward, P. S.  2005.  A synoptic review of the ants of California (Hymenoptera: Formicidae).  Zootaxa 936: 1-68. [PDF file]
  54. Ward, P. S., Brady, S. G.; Fisher, B. L.; Schultz, T. R. 2005. Assembling the ant “Tree of Life” (Hymenoptera: Formicidae). Myrmecologische Nachrichten 7: 87-90. [PDF file]
  55. Boulton, A. M., Davies, K. F., Ward, P. S. 2005. Species richness, abundance, and composition of ground-dwelling ants in northern California grasslands: role of plants, soil, and grazing. Environmental Entomology 34: 96-104. [PDF file]
  56. Brady, S. G., Ward, P. S. 2005. Morphological phylogeny of army ants and other dorylomorphs (Hymenoptera: Formicidae). Systematic Entomology 30: 593-618. [PDF file]
  57. van der Hammen, T.; Ward, P. S. 2005. Ants from the Ecoandes expeditions: diversity and distribution. Studies on Tropical Andean Ecosystems 6: 239-248. [PDF file]
  58. Suarez, A. V., Holway, D. A., Ward, P. S. 2005. The role of opportunity in the unintentional introduction of nonnative ants. Proceedings of the National Academy of Sciences of the United States of America 102: 17032-17035. [PDF file]
  59. Ward, P. S. 2006. The ant genus Tetraponera in the Afrotropical region: synopsis of species groups and revision of the T. ambigua-group (Hymenoptera: Formicidae). Myrmecologische Nachrichten 8:119-130. [PDF file]
  60. Ward, P. S.  2006.  Ants.  Current Biology 16: 152-155. [PDF file]
  61. Brady, S. G., Schultz, T. R., Fisher, B. L., Ward, P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172-18177. [PDF file]
  62. Bolton, B., Alpert, G., Ward, P. S., Naskrecki, P. 2007. Bolton’s Catalogue of ants of the world: 1758-2005. Cambridge, Mass.: Harvard University Press, CD-ROM.
  63. Ward, P. S. 2007. Edward O. Wilson and his contributions to ant systematics. Memoirs of the American Entomological Institute 80: 3-7. [PDF file]
  64. Ward, P. S. 2007. The ant genus Leptanilloides: discovery of the male and evaluation of phylogenetic relationships based on DNA sequence data. Memoirs of the American Entomological Institute 80: 637-649. [PDF file]
  65. Ward, P. S. 2007. Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae). Zootaxa 1668: 549-563. [PDF file]
  66. Ward, P. S., Brady, S. G. 2009. Rediscovery of the ant genus Amyrmex Kusnezov (Hymenoptera: Formicidae) and its transfer from Dolichoderinae to Leptanilloidinae. Zootaxa 2063: 46-54. [PDF file]
  67. Kautz, S., Lumbsch, H. T., Ward, P. S., Heil, M. 2009. How to prevent cheating: a digestive specialization ties mutualistic plant-ants to their ant-plant partners. Evolution 63: 839-853. [PDF file]
  68. Wernegreen, J. J., Kauppinen, S. N., Brady, S. G., Ward, P. S. 2009. One nutritional symbiosis begat another: phylogenetic evidence that the ant tribe Camponotini acquired Blochmannia by tending sap-feeding insects. BMC Evolutionary Biology (doi:10.1186/1471-2148-9-292) 9: Article 292. [PDF file]
  69. Ward, P. S. 2009. The ant genus Tetraponera in the Afrotropical region: the T. grandidieri group (Hymenoptera: Formicidae). Journal of Hymenoptera Research 18: 285-304. [PDF file]
  70. Ward, P. S. 2009. Taxonomy, phylogenetics, and evolution. Pp. 3-17 in: Lach, L., Parr, C. L., Abbott, K. (eds) 2009. Ant ecology. Oxford: Oxford University Press, xvii + 410 pp. [PDF file]
  71. Ward, P. S., Brady, S. G., Fisher, B. L., Schultz, T. R. 2010. Phylogeny and biogeography of dolichoderine ants: effects of data partitioning and relict taxa on historical inference. Systematic Biology 59: 342-362. [PDF file]
  72. Lucky, A., Ward, P. S. 2010. Taxonomic revision of the ant genus Leptomyrmex Mayr (Hymenoptera: Formicidae). Zootaxa 2688: 1-67. [PDF file]
  73. Ward, P. S. 2011. Integrating molecular phylogenetic results into ant taxonomy (Hymenoptera: Formicidae). Myrmecological News 15: 21-29. [PDF file]
  74. 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. A108: 220-225. [PDF file]
  75. 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 17: 5-11. [PDF file]
  76. Seltmann, K., Yoder, M., Miko, I., Forshage, M., Bertone, M., Agosti, D., Austin, A., Balhoff, J., Borowiec, M., Brady, S., Broad, G., Brothers, D., Burks, R., Buffington, M., Campbell, H., Dew, K., Ernst, A., Fernandez-Triana, J., Gates, M., Gibson, G., Jennings, J., Johnson, N., Karlsson, D., Kawada, R., Krogmann, L., Kula, R., Ohl, M., Rasmussen, C., Ronquist, F., Schulmeister, S., Sharkey, M., Talamas, E., Tucker, E., Vilhelmsen, L., Ward, P., Wharton, R., Deans, A. 2012. A hymenopterists’ guide to the Hymenoptera Anatomy Ontology: utility, clarification, and future directions. Journal of Hymenoptera Research 27: 67-88. [PDF file]
  77. Johnson, B. R., Borowiec, M. L., Chiu, J. C., Lee, E. K., Atallah, J., Ward, P. S. 2013. Phylogenomics resolves evolutionary relationships among ants, bees, and wasps. Current Biology 23: 2058-2062. [PDF file]
  78. 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. [PDF file]
  79. Ward, P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 23-43. [PDF file]
  80. 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. [PDF file]
  81. Ward, P. S., Brady, S. G., Fisher, B. L., Schultz, T. R. 2015. The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40: 61-81. [PDF file]
  82. 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. [PDF file]
  83. Ward, P. S., Blaimer, B. B., Fisher, B. L. 2016. A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and DinomyrmexZootaxa 4072: 343-357. [PDF file]
  84. Boudinot, B. E., Probst, R. S., Brandão, C. R. F., Feitosa, R. M., Ward, P. S. 2016. Out of the Neotropics: newly discovered relictual species sheds light on the biogeographic history of spider ants (Leptomyrmex, Dolichoderinae, Formicidae). Systematic Entomology 41: 658-671. doi:10.1111/syen.12181 [PDF file]
  85. Ward, P. S., Fisher, B. L. 2016. Tales of dracula ants: the evolutionary history of the ant subfamily Amblyoponinae (Hymenoptera: Formicidae). Systematic Entomology 41: 683-693. doi:10.1111/syen.12186 [PDF file]
  86. Ward, P. S., Brady, S. G., Fisher, B. L., Schultz, T. R. 2016. Phylogenetic classifications are informative, stable and pragmatic: the case for monophyletic taxa. Insectes Sociaux 63: 489-492 doi: 10.1007/s00040-016-0516-9 [PDF file]
  87. Ward, P. S. 2017. A review of the Pseudomyrmex ferrugineus and Pseudomyrmex goeldii species groups: acacia-ants and their relatives (Hymenoptera: Formicidae). Zootaxa, in press.
  88. Ward, P. S., Branstetter, M. G. 2017. The acacia ants revisited: convergent evolution and biogeographic context in an iconic ant/plant mutualism. Proceedings of the Royal Society B 284: 20162569. doi: 10.1098/rspb.2016.2569
  89. Branstetter, M. G., Longino, J. T., Ward, P. S., Faircloth, B. C. 2017. Enriching the ant tree of life: enhanced UCE bait set for genome-scale phylogenetics of ants and other Hymenoptera. Methods in Ecology and Evolution doi: 10.1111/2041-210X.12742
  90. Ward, P. S. 2017b. Subfamilia Pseudomyrmecinae. In: Fernández, F., Guerrero, R. & Delsinne, T. (Eds.), Hormigas de Colombia. Universidad Nacional de Colombia, Bogotá, in press.

Abstracts, Reviews

  • Ward, P. S. 1982. Colony structure and genetic relatedness in a species complex of ponerine ants. In Breed, M. D., Michener, C. D., Evans, H. E. (eds.) The biology of social insects; proceedings of the Ninth Congress IUSSI. Boulder: Westview Press, pp. 413-414 [Abstract].
  • Ward, P. S. 1983. Sex-ratio interactions between queenright and queenless colonies in rainforest ants of the Rhytidoponera impressa group. In Proceedings 18th Intern. Ethol. Conference, Brisbane, Qld., p. 299 [Abstract].
  • Ward, P. S. 1985. Review of: Wehner, R. et al. Foraging strategies in individually searching ants, Cataglyphis bicolorAnimal Behaviour 33:336-337.
  • Ward, P. S. 1995. Ants on the run. [Review of Gotwald, W. H., Jr. 1995. Army ants. The biology of social predation. Ithaca, New York: Cornell University Press, xviii + 302 pp.] Science 270:319-320.
  • Ward, P. S. 1998. Phylogeny and biogeography of host plant associations in Neotropical pseudomyrmecine ants (Hymenoptera: Formicidae). P. 499 in: Schwarz, M. P., Hogendoorn, K. (eds.) Social insects at the turn of the millenium. Proceedings of the XIII International Congress of IUSSI. Adelaide, Australia. 29 December 1998 – 3 January 1999. Adelaide: XIII Congress of IUSSI, 535 pp. [Abstract].
  • Brady, S. G., Gadau, J., Ward, P. S. 1999. Is the ant genus Camponotus paraphyletic? 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. [Abstract].
  • Ward, P. S.; Brady, S. G.  2002.  Phylogeny and biogeography of the ant genera Myrmecia and Nothomyrmecia (Hymenoptera: Formicidae) and related fossil taxa.  P. 156 in: XIV International Congress of IUSSI. The Golden Jubilee Proceedings. 27 July – 3 August 2002, Hokkaido University, Sapporo, Japan.  Sapporo: XIV International Congress of IUSSI, 233 pp. [Abstract.].

Limited Distribution

  • Sugden, E. A., Ward, P. S., Shapiro, A. M., Teague, S. 1985. Invertebrates. Pp. 47-62 in: Weathers, W. W., Cole, R. (eds.) Flora and fauna of the Stebbins Cold Canyon Reserve, Solano County, California. Institute of Ecology Publication No. 29. University of California, Davis. vi + 84 pp.
  • Ward, P. S. 1990. The endangered ants of Mauritius: doomed like the dodo? Notes from Underground No. 4 (Spring 1990): 3-5.
  • Ward, P. S. 1994. Les fourmis. P. 30 in: Goodman, S. N., Langrand, O. (eds.) Inventaire biologique Forêt de Zombitse. Recherches pour le Développement. Série Sciences Biologiques. No. Spécial. Antananarivo, Madagascar: Centre d’Information et de Documentation Scientifique et Technique, 106 pp.