My research explores the interactions of food (prey) quality and quantity on consumer dynamics and thence on trophic dynamics (stoichiometric ecology) and biogeochemistry. Targets for this work are primarily planktonic and microbial systems, with recent emphasis upon mixotrophic plankton and variations in predation rates with prey size, abundance and nutritional quality. Applied aspects include optimisation of algal biomass production in the presence of zooplanktonic pests, and microbial activity in wetlands. The tools for research are primarily dynamic adaptive multi-nutrient (variable stoichiometry) models. I am currently involved in various EU, RCUK and Welsh funded projects investigating a variety of issues ranging from sustainable use of water to the impact of human behaviour on the global oceans. I have also experienced life on the other side of research as Managing Editor of the Journal of Plankton Research (a peer-reviewed scientific journal publication from Oxford University Press), a climate research consultant for the Welsh Local Government Association, and as Biodiversity Officer for Bridgend County Borough Council.

Areas of Expertise

  • Systems Dynamics Modeller
  • PhD Zooplankton Growth Dynamics; A Modelling Study (SAMS/Open University, UK)
  • MSc Ecosystems Analysis & Governance (Warwick; UK)
  • BSc (Hons) Botany (Presidency, India) 1st Class


  1. & Sampling bias misrepresents the biogeographical significance of constitutive mixotrophs across global oceans. Global Ecology and Biogeography 28(4), 418-428.
  2. & Exploring nonlinear functional responses of zooplankton grazers in dilution experiments via optimization techniques. Limnology and Oceanography 64(2), 774-784.
  3. The Perfect Beast. Scientific American 318(4), 26-33.
  4. & Modelling mixotrophic functional diversity and implications for ecosystem function. Journal of Plankton Research 40(6), 627-642.
  5. & Simulating Effects of Variable Stoichiometry and Temperature on Mixotrophy in the Harmful Dinoflagellate Karlodinium veneficum. Frontiers in Marine Science 5
  6. & Toward a mechanistic understanding of trophic structure: inferences from simulating stable isotope ratios. Marine Biology 165(9)
  7. & Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance. Proceedings of the Royal Society B: Biological Sciences 284(1860), 20170664
  8. & Minimising losses to predation during microalgae cultivation. Journal of Applied Phycology 29(4), 1829-1840.
  9. & Modeling Plankton Mixotrophy: A Mechanistic Model Consistent with the Shuter-Type Biochemical Approach. Frontiers in Ecology and Evolution 5
  10. & Introducing mixotrophy into a biogeochemical model describing an eutrophied coastal ecosystem: The Southern North Sea. Progress in Oceanography 157, 1-11.
  11. & Biological or microbial carbon pump? The role of phytoplankton stoichiometry in ocean carbon sequestration. Journal of Plankton Research
  12. et. al. Defining Planktonic Protist Functional Groups on Mechanisms for Energy and Nutrient Acquisition: Incorporation of Diverse Mixotrophic Strategies. Protist 167(2), 106-120.
  13. & Mixotrophy in the Marine Plankton. Annual Review of Marine Science 9(1), 311-335.
  14. & Exploring the Implications of the Stoichiometric Modulation of Planktonic Predation. In Aquatic Microbial Ecology and Biogeochemistry: A Dual Perspective. (pp. 77-89). Springer International Publishing.
  15. & Why Plankton Modelers Should Reconsider Using Rectangular Hyperbolic (Michaelis-Menten, Monod) Descriptions of Predator-Prey Interactions. Frontiers in Marine Science 3
  16. & The role of mixotrophic protists in the biological carbon pump. Biogeosciences 11(4), 995-1005.
  17. & Bridging the gap between marine biogeochemical and fisheries sciences; configuring the zooplankton link. Progress in Oceanography 129, 176-199.
  18. & Decrease in diatom palatability contributes to bloom formation in the Western English Channel. Progress in Oceanography 137, 484-497.
  19. & Ocean acidification with (de)eutrophication will alter future phytoplankton growth and succession. Proceedings of the Royal Society B: Biological Sciences 282(1804), 20142604-20142604.
  20. & Impact of zooplankton food selectivity on plankton dynamics and nutrient cycling. Journal of Plankton Research 37(3), 519-529.
  21. & Misuse of the phytoplankton-zooplankton dichotomy: the need to assign organisms as mixotrophs within plankton functional types. Journal of Plankton Research 35(1), 3-11.
  22. & Acclimation, adaptation, traits and trade-offs in plankton functional type models: reconciling terminology for biology and modelling. Journal of Plankton Research 37(4), 683-691.
  23. & Prymnesium parvum invasion success into coastal bays of the Gulf of Mexico: Galveston Bay case study. Harmful Algae 43, 31-45.
  24. et. al. Mechanisms of microbial carbon sequestration in the ocean – future research directions. Biogeosciences 11(19), 5285-5306.
  25. & Introduction to the BASIN Special Issue: State of art, past present a view to the future. Progress in Oceanography 129, 171-175.

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  • BIO017 Key Skills for Biologists

    This module is designed to guide students through the fundamental tools needed to successfully review their timetable and coursework deadlines, submit coursework, check rules and regulations, and develop important communication, teamwork and problem-solving skills. Through ICT workshops, group work and laboratory work, and in partnership with tutorials, this module covers key elements required for students' undergraduate studies and subsequently as a professional scientist.

  • BIO227 Marine Plankton And Oceanography

    This module introduces students to the fundamental concept of plankton ecology at level 2. Students will receive 18-20 lectures and four practicals (1 wet lab, 3 IT lab, 1 boat work). The lectures will cover three key themes: oceanography pertaining to planktonic production, plankton ecophysiology, and the functioning of the planktonic foodwebs. Three reports (1 IT-based, 1 from a lab-practical, 1 from boat work) will be assessed, together with one examination consisting of 30 multiple choice questions + one essay question + one analytical question.


  • Accounting for mixotrophy in marine microbial food webs (current)

    Student name:
    Other supervisor: Prof Kevin Flynn