Professor of Mathematics,
Mathematical modeling and statistical analyses of infectious diseases

NCSU Statistics Department
2311 Stinson Drive
Campus Box 8203
Raleigh, NC 27695-8203

Office: 308 Cox Hall, North Carolina State University
Phone: 919 515-1910
Mail: Campus Box 8213, 308 Cox Hall,
North Carolina State University,
Raleigh, NC 27695-8213


I studied mathematics at Trinity College, Cambridge, before moving to the Department of Zoology in Oxford to do a Ph.D. with Robert May, which I completed in 1996. The following three years in Oxford saw me doing my first postdoc, on a Medical Research Council Non-Clinical Fellowship, and a lectureship at St. Hilda’s College. In 1999 I moved to the US, for a four year stint as a Long-Term Member in the Institute for Advanced Study‘s Program in Theoretical Biology. I moved to my NC State faculty position in 2003.



I am a mathematical biologist. The majority of my work concerns the epidemiology of infectious diseases, with a particular recent interest in mosquito-borne infections.  I am a professor in the Department of Mathematics at North Carolina State University, I also direct the Biomathematics Graduate Program and the Research Training Group in Mathematical Biology.



  1. Lessler, J., Edmunds, W.J., Halloran, M.E., Hollingsworth, T.D. & Lloyd, A.L. (2014). Seven challenges for model-driven data collection in experimental and observational studies. Epidemics. In press. PDFdoi:10.1016/j.epidem.2014.12.002
  2. Roberts, M., Andreasen, V., Lloyd, A. & Pellis, L. (2014). Nine challenges for deterministic epidemic models. Epidemics. In press. doi:10.1016/j.epidem.2014.09.006 PDF
  3. Hollingsworth, T.D., Pulliam, J.R.C., Funk, S., Truscott, J.E., Isham, V. & Lloyd, A.L. (2014). Seven challenges for modelling indirect transmission: vector-borne diseases, macroparasites and neglected tropical diseases. Epidemics. In press. doi:10.1016/j.epidem.2014.08.007 PDF
  4. Britton, T., House, T., Lloyd, A.L., Mollison, D., Riley, S. & Trapman, P. (2014). Eight challenges for stochastic epidemic models involving global transmission. Epidemics. In press. PDF
  5. Robert, M.A., Okamoto, K.W., Gould, F. & Lloyd, A.L. (2014). Anti-pathogen genes and the replacement of disease-vectoring mosquito populations: a model-based evaluation. Evol. Applic. 7:1238–1251. doi:10.1111/eva.12219 PDF Info.
  6. Guerra, C.A., Reiner, R.C., Perkins, T.A., Lindsay, S.W., Midega, J., Brady, O.J., Barker, C.M., Reisen, W.K., Harrington, L.C., Taken, W., Kitron, U., Lloyd, A.L., Hay, S.I., Scott, T.W. & Smith, D.L. (2014). An assembly of adult female mosquito mark-release-recapture data to inform vector-borne pathogen transmission models. Parasites and Vectors. 7:276. doi:10.1186/1756-3305-7-276 PDF
  7. Reiner, R.C., Stoddard, S.T., Forshey, B.M., King, A.A., Ellis, A.M., Lloyd, A.L., Long, K.C., Rocha, C., Vilcarromero, S., Astete, H, Bazan, I., Lenhart, A., Vazquez-Prokopec, G.M., Paz-Soldan, V.A., McCall, P.J., Kitron, U., Elder, J.P., Halsey, E.S., Morrison, A., Kochel, T.J., Scott. T.W. (2014). Time-varying, serotype-specific force of infection estimates for dengue virus using 12 years of longitudinal serological data from Iquitos, Peru. Proc. Natl. Acad. Sci.111: E2694-E2702. doi:10.1073/pnas.1314933111 PDFPDF with Supp. Info.
  8. Okamoto, K.W., Robert, M.A., Gould, F. & Lloyd, A.L. (2014). Feasible introgression of an anti-pathogen transgene into an urban mosquito population without using gene-drive. PLoS Negl. Trop. Dis. 8(7): e2827. doi:10.1371/journal.pntd.0002827 PDF Info.
  9. Smith, D.L., Perkins, T.A., Reiner, R.C., Barker, C.M., Niu, T., Chaves, L.F., Ellis, A.M., George, D.B., Le Menach, A., Pulliam, J.R.C., Bisanzio, D., Buckee, C., Chiyaka, C., Cummings, D.A.T., Garcia, A.J., Gatton, M.L., Gething, P.W., Hartley, D.M., Johnston, G., Klein, E.Y., Michael, E., Lloyd, A.L., Piggot, D.M., Reisen, W.K., Ruktanonchai, N., Singh, B.K., Stoller, J., Tatem, A.J., Kitron, U., Godfray, H.C.J., Cohen, J.M., Hay, S.I. & Scott, T.W. (2014). Recasting the Theory of Mosquito-Borne Pathogen Transmission Dynamics and Control. Trans. R. Soc. Trop. Med. Hyg. 108, 185-197. DOI:10.1093/trstmh/tru026 PDF
  10. Yarmand, H., Ivy, J.S., Denton, B. & Lloyd, A.L. (2014). Optimal Two-Phase Vaccine Allocation to Geographically Different Regions under Uncertainty. Eur. J. Oper. Res. 233, 208-219.
  11. Okamoto, K.W., Robert, M.A., Lloyd, A.L. & Gould, F. (2013). A Reduce and Replace strategy for suppressing vector-borne diseases: Insights from a stochastic, spatial model. PLoS ONE 8(12):e81860. DOI:10.1371/journal.pone.0081860 PDF.
  12. Legros, M., Xu, C., Morrison, A., Scott, T.W., Lloyd, A.L. & Gould, F. (2013). Modeling the dynamics of a non-limited and a self-limited gene drive system in structured Aedes aegypti PLoS ONE 8: e83354. DOI:10.1371/journal.pone.0083354 PDF
  13. Scholle, S.O., Ypma, R.J.F., Lloyd, A.L. & Koelle, K. (2013). Viral substitution rate variation can arise from the interplay between within-host and epidemiological dynamics. Am. Nat.182, 494-513.
  14. Robert, M.A., Okamoto, K., Lloyd, A.L. & Gould, F. (2013). A Reduce and Replace strategy for suppressing vector-borne diseases: Insights from a deterministic model. PLoS ONE8(9): e73233. DOI:10.1371/journal.pone.0073233 PDF
  15. Walsh, R.K., Aguilar, C.L., Facchinelli, L., Valerio, L., Ramsey, J.M., Scott, T.W., Lloyd, A.L. & Gould, F. (2013). Regulation of Aedes aegyptipopulation dynamics in field systems: Quantifying direct and delayed density dependence. Am. J. Trop. Med. Hyg. 89, 68-77. DOI:10.4269/ajtmh.12-0378
  16. Breen, M., Villeneuve, D.L., Ankley, G.T., Bencic, D.C., Breen, M.S., Watanabe, K.H., Lloyd, A.L. & Conolly, R.B. (2013). Developing Predictive Approaches to Characterize Adaptive Responses of the Reproductive Endocrine Axis to Aromatase Inhibition: II. Computational Modeling. Toxicol. Sci. 133, 234-247.
  17. Reiner, R.C., Perkins, T.A., Barker, C.M., Niu, T., Chaves, L.F., Ellis, A.M., George, D.B., Le Menach, A., Pulliam, J.R.C., Bisanzio, D., Buckee, C., Chiyaka, C., Cummings, D.A.T., Garcia, A.J., Gatton, M.L., Gething, P.W., Hartley, D.M., Johnston, G., Klein, E.Y., Michael, E., Lindsay, S.W., Lloyd, A.L., Piggot, D.M., Reisen, W.K., Ruktanonchai, N., Singh, B.K., Tatem, A.J., Kitron, U., Hay, S.I., Scott, T.W., & Smith, D.L. (2013). A systematic review of mathematical models of mosquito-borne pathogen transmission: 1970-2010. J. R. Soc. Interface. 10: 20120291. PDF
  18. Facchinelli, L., Valerio, L., Ramsey, J.M., Gould, F., Walsh, R.K., Bond, G., Robert, M.A., Lloyd, A.L., James, A.A., Alphey, L. & Scott, T.W. (2013). Field cage studies and progressive evaluation of genetically-engineered mosquitoes. PLoS Negl. Trop. Dis. 7(1): e2001. PDF
  19. Legros, M., Xu, C., Okamoto, K., Scott, T.W., Morrison, A.C., Lloyd, A.L. & Gould, F. (2012). Assessing the feasibility of controlling Aedes aegyptiwith transgenic methods: A model-based evaluation. PLoS ONE 7(12): e52235. PDF
  20. Robert, M.A., Legros, M., Facchinelli, L., Valerio, L., Ramsey, J.M., Scott, T.W., Gould, F. & Lloyd, A.L. (2012). Mathematical models as aids for design and interpretation of experiments: The case of transgenic mosquitoes. J. Med. Entomol. 49, 1177-1188.
    PDFSupplementary information: Appendix 1, Appendix 2
  21. Capaldi, A., Behrend, S., Berman, B., Smith, J. & Lloyd, A.L. (2012). Parameter estimation and uncertainty quantification for an epidemic model. Math. Biosci. Eng. 9, 553-576. PDF
  22. Durrett, R., Gleeson, J.P., Lloyd, A.L., Mucha, P.J., Shi, F., Sivakoff, D., Socolar, J. & Varghese, C. (2012). Graph fission in an evolving voter model. Proc. Natl. Acad. Sci. USA. 109, 3682-3687. PDF
  23. Walker, T., Johnson, P.H., Moreira, L.A., Iturbe-Ormaetxe, I., Frentiu, F.D., McMeniman, C.J., Leong, Y.S., Dong, Y., Axford, J., Kriesner, P., Lloyd, A.L, Ritchie, S.A., O’Neill, S.L. & Hoffmann, A.A. (2011). The wMel Wolbachiastrain blocks dengue and invades caged Aedes aegypti Nature 456, 450-453.
  24. Huang, Y., Lloyd, A.L., Legros, M. & Gould, F. (2011). Gene-drive into insect populations with age and spatial structure: A theoretical assessment. Evol. Applic. 4(3), 415-428.doi:10.1111/j.1752-4571.2010.00153.x
  25. Breen, M., Breen, M.S., Terasaki, N. Yamazaki, M., Lloyd, A.L. & Conolly, R.B. (2011). Mechanistic computational model of steroidogenesis in H295R cells: Role of oxysterols and cell proliferation to improve predictability of biochemical response to endocrine active chemicals. Toxicol. Sci. 123(1), 80-93.
  26. Legros, M., Magori, K., Morrison, A.C., Xu, C., Scott, T.W., Lloyd, A.L. & Gould, F. (2011). Evaluation of location-specific predictions by a detailed simulation model of Aedes aegypti PLoS ONE 6(7), e22701. doi:10.1371/journal.pone.0022701 PDF
  27. Ward, C.M., Su, J.T., Huang, Y., Lloyd, A.L., Gould, F. & Hay, B.A. (2011). Medeaselfish genetic elements as tools for altering traits of wild populations: A theoretical analysis. Evolution. 65(4), 1149-1162. doi:10.1111/j.1558-5646.2010.01186.x
  28. Xu, C., Legros, M., Gould, F. & Lloyd, A.L. (2010). Understanding uncertainties in model-based predictions of the population dynamics of Aedes aegypti. PLoS Negl. Trop. Dis. 4(9), e830. doi:10.1371/journal.pntd.0000830 PDFSupplementary information
  29. Arauz, L.F., Neufeld, K.N., Lloyd, A.L. & Ojiambo, P.S. (2010). Quantitative models for germination and infection of Psuedoperonospora cubensis in response to temperature and duration of leaf wetness. Phytopathology 100, 959-967.