Professor John Spicer

Professor John Spicer


School of Marine Science and Engineering (Faculty of Science and Engineering)


Professor of Marine Zoology


B.Sc.(Hons) Zoology, Glasgow 1982

Ph.D. in Zoology, Glasgow 1987

P.G.C.E.(S) (with distinctions), Jordanhill College

Glasgow 1991

Authored books

Spicer JI: Biodiversity: A beginner's guide. Oneworld Books, Oxford (2006); World Issues Today, Ransom Publishers (2009); Audiobook (2012)

Gaston KJ, Spicer JI: Biodiversity: An introduction. Blackwells 2nd edition (2003), 1st edition (1998). Translated into Greek, Spanish & Chinese. 

Barnes RSK, Calow P, Olive PJW, Golding DW, Spicer JI: The invertebrates: a new synthesis (3rd ed). Blackwell Scientific, Oxford (2001). Translated into Spanish and Chinese. 

Spicer JI, Gaston KJ: Physiological diversity and its ecological implications. Blackwells Science, Oxford (1999). Translated into Chinese. 


Scientific Publications

In press

Bitterli, T., Rundle, S.D. and J.I. Spicer: Development of cardiovascular function in the marine gastropod Littorina obtusata (Linnaeus). Journal of Experimental Biology, in press.

De La Haye, K., Spicer, J.I., Widdicombe, S. and M. Briffa: Reduced pH sea water disrupts chemo-responsive behaviour in an intertidal crustacean. Journal of Experimental Biology and Ecology. Available online 10.1016/j.jembe.2011.11.013


Bozinovic, F., Calosi, P. and J.I. Spicer: Physiological correlates of geographical range in animals. Annual Review of Ecology, Evolution and Systematics Volume 42, pages 155 – 179 (2011)

Calosi, P., Bilton, D.T., Spicer, J.I., Verberk, W.C.E.P., Atfield, A. and T. Garland: The comparative biology of diving in two genera of European Dytiscidae. Journal of Evolutionary Physiology, Volume 25, pages 329 – 341 (2012)

de la Haye, K.L., Spicer, J.I., Widdicombe, S. and M. Briffa: Low pH disrupts resource assessment and decision making in the hermit crab Pagurus bernhardus. Animal Behaviour, Volume 82, pages 495 – 501 (2011)

Ellis, R.P., Parry, H., Spicer, J.I., Hutchinson, T.H., Pipe, R.K. and S. Widdicombe: Immunological function in marine invertebrates: Responses to environmental perturbation. Fish and Shellfish Immunology, Volume 30, pages 1209 – 1222 (2011)

Findlay, H.S., Wood, H.L., Kendall, M.A., Spicer, J.I., Twitchett, R.  and S. Widdicombe: Comparing the impact of high CO2 on calcium carbonate structures in different marine organisms. Marine Biology Research. Volume 7, pages 565 – 575 (2011)

Morley, S.A., Lemmon, V., Obermüller, B.E., Spicer, J.I., Clark, M.S. and L.S. Peck : Duration tenacity: a method for assessing acclimatory capacity of the Antarctic limpet, Nacella concinna. Journal of Experimental Biology and Ecology, Volume 399, pages 39 – 42 (2011)

Rundle, S.D., Smirthwaite, J.J., Colbert, M.W. and J.I. Spicer: Predation cues alter the timing of developmental events in gastropod embryos. Biology Letters, Volume 7, pages 285 – 287 (2011)

Spicer, J.I.: Darwin and God. Science & Christian Belief , Volume 23, pages 87 – 88 (2011)

Spicer, J.I., Rundle, S.D. and O. Tills:  Studying the altered timing of physiological events during development. It's about time... or is it? Respiratory Physiology & Neurobiology, Volume 178, pages 3 – 12 (2011)

Spicer, J.I., Widdicombe, S., Needham, H.R. and J.A. Berge: Impact of CO2-acidified  seawater on the extracellular acid-base balance of the northern sea urchin Strongylocentrotus dröebachiensis. Journal of Experimental Marine Biology and Ecology, Volume 407, pages 19 – 25 (2011)

Tills, O., Rundle, S.D., Salinger, M., Haun, T., Pfenninger, M. and J.I. Spicer: A genetic basis for intra-specific differences in developmental timing? Evolution and Development, Volume 13, pages 542 – 548 (2011)

Verberk, W.C.E.P., Bilton, D.T., Calosi, P. and J.I. Spicer: Limits to oxygen supply in aquatic ectotherms. Ecology, Volume 92, pages 1565 – 1572 (2011)

Widdicombe, S., Spicer, J.I. and V. Kitidis: Effects of ocean acidification on sediment fauna. Chapter 11. In, Ocean Acidification (eds Gattuso, J.P. and L. Hansson). Oxford University Press (2011).

Wood, H.L., Spicer, J.I., Kendall, M.A., Lowe, D.M. and S. Widdicombe: Ocean warming and acidification; implications for the Arctic brittlestar Ophiocten sericeum. Polar Biology, Volume 34, pages 1033 – 1044 (2011)


Calosi, P., Bilton, D.T., Spicer, J.I., Votier, S., and A. Atfield: What determines a species’ geographical range? Thermal biology and latitudinal range size relationships in European diving beetles (Coleoptera: Dytiscidae). Journal of Animal Ecology, Volume 79, pages 194 – 204 (2010).

Dissanayake, A., Clough, R., Spicer, J.I. and  M.B. Jones: Effects of hypercapnia on acid–base balance and osmo-/iono-regulation in prawns (Decapoda: Palaemonidae). Aquatic Biology, Volume 11, pages 27 - 36 (2010).

Findlay, H.S., Kendall, M.A., Spicer, J.I. and S. Widdicombe: Post-larval development of two intertidal barnacles at elevated CO2 and temperature. Marine Biology (Berlin)Volume 157, pages 725 – 735 (2010).

Findlay, H.S., Kendall, M.A., Spicer, J.I. and S Widdicombe: Relative influences of ocean acidification and temperature on intertidal barnacle post-larvae at the northern edge of their geographic distribution. Estuaries, Coastal and Shelf Science, Volume 86, pages 675-682 (2010).

Findlay, H.S., Burrows, M.T., Kendall, M.A, Spicer J.I. and S. Widdicombe: Can ocean acidification affect population dynamics of Semibalanus balanoides at its southern range edge. Ecology, Volume 91, pages 2931 – 2940 (2010).

Janas, U. and J.I. Spicer: Seasonal and temperature effects on osmoregulation by the invasive prawn Palaemon elegans Rathke, 1837 in the Baltic Sea. Marine Biology Research, Volume 6, pages 333 – 337 (2010).

Marchant, H., Calosi, P. and J.I. Spicer: Exposure to mild CO2-acidification of sea water does not compromise acid-base balance, metabolism or feeding rate of the limpet Patella vulgata but surprisingly is accompanied by radula damage.  Journal of the Marine Biological Association of the United Kingdom Volume 90, pages 1379 – 1384 (2010).

Mourabit, S., Rundle, S.D., Spicer, J.I. and K.A. Sloman:  Alarm substance from adult zebrafish alters early embryonic development in offspring. Biology Letters, Volume 6, pages 525 – 528 (2010).

Sánchez-Fernández, Calosi, P., Atfield, A., Arribas, P., Velasco, J., Spicer, J.I., Millán, A and D.T. Bilton: Reduced salinities compromise the thermal tolerance of hypersaline specialist diving beetles (Coleoptera, Dytiscidae). Physiological Entomology, Volume 35, pages 265 – 273 (2010).

Small, D., Calosi, P. White, D., Spicer, J.I. and S. Widdicombe: Impact of medium term exposure to CO2 enriched seawater on the physiological functions of the velvet swimming crab, Necora puber. Aquatic Biology Volume 10, pages 11 – 21 (2010).

Spicer, J.I. and R. Saborowski: Physiology and metabolism of Northern krill. Chapter 4. In, Biology of Northern Krill (ed. Tarling, G.). Advances in Marine Biology, Volume 57, pages 91 – 126 (2010).

Tills, O., Spicer, J.I. and S.D. Rundle: Salinity-induced heterokairy in an upper-estuarine population of the snail Radix balthica (Mollusca: Pulmonata). Aquatic Biology, Volume 9, pages 95 – 105 (2010).

Wood, H.L., Spicer, J.I., Lowe, D.M. and S. Widdicombe: Interaction of ocean acidification and temperature; the high cost of survival in the brittlestar Ophiura ophiura. Marine Biology (Berlin), Volume 157, pages 2001 – 2013 (2010).


Arnold, K.E., Findlay, H.S., Boothroyd, D., Daniels, C.L and J.I. Spicer: Effect of hypercapnia-related acidification on the larval development of the European lobster, Homarus gammarus (L.). Biogeosciences, Volume 6, pages 1747 – 1754 (2009)

Arnold, K.E., C. Wells and J.I. Spicer:  Effect of an insect juvenile hormone analogue, Fenoxycarb® on development and oxygen uptake by larval lobsters Homarus gammarus (L.). Comparative Biochemistry and Physiology C, Volume 149, pages 393 – 396 (2009)

Byrne, R.A., Rundle, S.D., Smirthwaite, J.J. and J.I. Spicer: Embryonic rotational behaviour in the pond snail Lymnaea stagnalis; influence of environmental oxygen and developmental stage. Zoology- Analysis of Complex Systems, Volume 112, pages 471 – 477 (2009)

Egilsdottir, H., Spicer, J.I. and S.D. Rundle: The effect of CO2 acidified sea water and reduced salinity on aspects of the embryonic development of the amphipod Echinogammarus marinus (Leach). Marine Pollution Bulletin. Volume 58, pages 1187 – 1191 (2009)

Ellis, R.P., J. Bersey, S.D. Rundle, J.M. Hall-Spencer and J.I. Spicer: Subtle but significant effects of CO2 acidified sea water on embryos of the intertidal snail Littorina obtusata. Aquatic Biology, Volume 5, pages 41 – 48 (2009).

Findlay, H.S., Kendall, M.A., Spicer, J.I. and S. Widdicombe: Future high CO2 in the intertidal may compromise adult barnacle Semibalanus balanoides survival and embryonic development rates. Marine Ecology – Progress Series, Volume 389, pages 193 – 202 (2009)

Findlay, H.S., Wood, H.L., Kendall, M.A., Spicer, J.I., Twitchett, R.J. and S. Widdicombe: Calcification, a physiological process to be considered in the context of the whole organism. Biogeosciences Discussions, Volume 6, pages 2267 – 2284 (2009)

Gaston, K.J., Chown, S.L., Calosi, P., Bernardo, J., Bilton, D.T., Clarke, A., Clusella-Trullas, S., Ghalambor, C.K., Konarewski, M., Peck, L.S., Porter, W.P., Pörtner, H.O., Rezende, E.L., Shulte, P.M., Spicer, J.I., Stillman, J., Terblanche, J.S. and M. van Kleunen: Macrophysiology: A conceptual re-unification. American Naturalist, Volume 174, pages 595 – 612 (2009)

Smirthwaite, J.J., S.D. Rundle and J.I. Spicer: The use of developmental sequences for assessing evolutionary change in gastropods. American Malacological Bulletin, Volume 27, pages 105 – 111 (2009)

Wood, H.L., Widdicombe, S. and J.I. Spicer: The influence of hypercapnia and macrofauna on sediment nutrient flux - will ocean acidification affect nutrient exchange? Biogeosciences Discussions, Volume 6, pages 2387 – 2413 (2009)


Bibby, R., S. Widdicombe, H. Parry, J.I. Spicer and R. Pipe.: Impact of ocean acidification on the immune response of the blue mussel, Mytilus edulis. Aquatic Biology. Volume 2, pages 67 - 74 (2008).

Calosi, P., D. Bilton and J.I. Spicer: Thermal tolerance, acclimatory capacity and vulnerability to global climate change. Biology Letters. Volume 4, 99 – 102 (2008).

Calosi, P., D. Bilton, J.I. Spicer, and A. Atfield: Thermal tolerance and geographic range size in the Agabus brunneus group of European diving beetles (Coleoptera: Dytiscidae). Journal of Biogeography. Volume 35, pages 295 - 305 (2008).

Findlay, H., M.A. Kendall, J.I. Spicer, C. Turley and S. Widdicombe: A novel microcosm system for investigating the impacts of elevated carbon dioxide and temperature on intertidal organisms. Aquatic Biology, Volume 3, pages 51 – 62 (2008).

Janas, U. and J.I. Spicer: Does the effect of low temperature on osmoregulation by the prawn Palaemon elegans Rathke, 1837 explain    winter migration offshore? Marine Biology (Berlin). Volume 153, pages 937 – 943 (2008).

Mowles, S., M. Briffa, P. Cotton and J.I. Spicer: The role of circulating metal ions during shell fights in the hermit crab Pagurus bernhardus. Ethology 114 1014 – 1022 (2008).

Widdicombe, S. and J.I. Spicer: Predicting the impact of ocean acidification on benthic biodiversity: What can physiology tell us? Journal of Experimental Marine Biology and Ecology Volume 366, pages 187 – 197 (2008).

Wood, H., S. Widdicombe and J.I. Spicer: Ocean Acidification may increase calcification rates- but at a cost. Proceedings of the Royal Society, London, Series B., Volume 275, pages 1767 – 1773 (2008).


Bibby, R. P. Cleall-Harding, S. Rundle, S. Widdicombe. and J. Spicer: Ocean acidification disrupts induced defences in the intertidal gastropod Littorina littorea. Biology Letters. Volume 3, pages 699 - 701 (2007).

Calosi, P., D. Bilton and J.I. Spicer:  The diving response of the diving beetle Ilybius montanus (Coleoptera: Dytiscidae): the effects of temperature and acidification. Journal of Zoology. Volume 273, pages 289 – 297 (2007).

Miles, H., S. Widdicombe, J.I. Spicer and J. Hall-Spencer: Effects of anthropogenic seawater acidification on acid-base balance in the sea urchin Psammechinus miliaris. Marine Pollution Bulletin. Volume 54, 89 – 96 (2007). Sanders, J., M. Kendell, A.J.S. Hawkins and J.I. Spicer: Can functional groups be used to indicate estuarine ecological status. Hydrobiologia. Volume 588, pages 45 – 58 (2007). Smirthwaite, J., S.D. Rundle, O.R.P. Bininda-Emonds and J.I. Spicer: An integrative approach identifies sequence heterochronies in pulmonate snails. Evolution and Development. Volume 9, pages 122 - 130 (2007). Spicer, J.I., A. Raffo and S. Widdicombe: Influence of CO2-related sea water acidification on extracellular acid-base balance in velvet fiddler crab Necora puber. Marine Biology (Berlin). Volume 151, pages 1117 - 1125 (2007).

Spicer, J.I. and S.D. Rundle: Plasticity in the timing of physiological development: Physiological heterokairy - what is it, how frequent is it, and does it matter? Comparative Biochemistry and Physiology, Part A. Volume 148, pages 712 - 719 (2007).