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Undergraduate courses

Dr Lewis Halsey

Dr Lewis Halsey

Senior Lecturer in Comparative and Environmental Physiology

Department of Life Sciences


My body of publications in the main covers four topics, all of which primarily concern vertebrate environmental physiology and energetics: the respiratory physiology, energetics and behaviour of ducks and cormorants; the relationships between the behaviour, ecology and energetics of wild diving king penguins; comparative analysis of diving and pedestrian locomotion across species, and the development of the 'accelerometry technique' as a method for quantifying behaviour and energy expenditure in terrestrial and aquatic animals. In the past I have also written occasional papers on subjects outside of this field such as on facial asymmetry perception and the high noise to signal ratios of football matches. 

A full list of my publications is available here - GoogleScholar and ResearcherID - access to the majority of them as pdfs can be found here - ResearchGate - and here -

Research interests

My present research takes a comparative approach in seeking to quantify the energetic costs for species in situations where energy expenditure can be affected and have marked consequences for fitness. For example, the comparative costs of terrestrial locomotion in animals; putative changes in the energy expenditure of people dependent on whether they fast through the morning or not; the calories expended by king penguins while walking; the energy costs for primates of using different methods of arboreal locomotion; associations between reproductive success and foraging energetic efficiency in seabirds.

Fundamental to such questions is trying to understand the variability in behavioural and physiological adaptations of animals, including humans, in two respects. First, in terms of adaptive plasticity to a changing environment, where habitat changes occur both due to seasonal fluctuations, and due to year on year variations as a result of human influences. Second, in terms of variations in adaptiveness and perhaps ultimately, therefore, reproductive success between individuals within a population. Behavioural, physiological and energetic measures in combination enable me to answer fundamental questions about the plasticity of animals and humans.

Particularly for fieldwork but also for laboratory based studies, I consider the application of miniaturised data loggers to record key behavioural and physiological variables to be a powerful technique. I employ and develop, in particular, the use of acceleration loggers for recording both the behaviours and associated energetics of wild animals, mainly in collaboration with the groups led by:

Dr Susannah Thorpe, University of Birmingham

Dr Charles Bishop, Bangor University

Dr Jon Green, University of Liverpool

Dr Yves Handrich, CNRS Strasbourg

Prof. Rory Wilson, Swansea University

Dr Craig White, University of Queensland

Dr Shaun Killen, University of Glasgow

Research projects undertaken

I am presently involved in several on-going funded projects including:

- Studying the behaviour, movements, macro-physiology and thus developing conservation management plans for seabird species inhabiting the Caribbean including the British Virgin Islands, funded by DEFRA Darwin Plus and the Leverhulme Trust (

- Studying the costs of arboreal locomotion in tree-dwelling primates (NERC funded).

- Assessing the validity of using heart rate loggers and acceleration loggers to measure the costs of wing adornments in volant birds (Royal Society funded).

- On going development of a seabird field site on Puffin Island in Wales, primarily to study intra-specific variation in species exhibiting reproductive skew (Research at Puffin Island; internally funded - Roehampton and Liverpool).

- Investigating aspects of variation in the behaviour and physiology of breakfast consumption (Industrial funding).

Present recent research associates and Ph. D students, and alumni 

Yvonne McMeel, externally funded.
Differences between breakfast eaters and breakfast skippers as a result of morningness and eveningness preferences and sleep habits.

Dr Tina Smith, externally funded.
Uncovering the mechanisms explaining differences in body mass index between breakfast eaters and breakfast skippers. 

Dr Sam Coward, NERC funded.
Energy costs and savings of arboreal locomotion in great apes: measuring a tractable model species - humans.

Astrid Willener, (PhD student), funded by the University of Roehampton and CNRS, France.
The biomechanics and energetics of a common behaviour for poorly-adapted species: a thorough analysis of pedestrian locomotion in penguins. (Field work undertaken on the Crozet Archipelago). Co-supervisors: Dr Yves Handrich, CNRS Strasbourg; Dr Siobhan Strike, University of Roehampton.

Gillian Lyons, (PhD student), funded by Queen's University of Belfast. (Secondary Supervisor)
Aspects of the behavioural and physiological ecology of the cuttlefish Sepia officinalis with relation to present and future oceans. Primary supervisor: Dr Jon Houghton, Queen's University Belfast; Co-supervisor: Dr Hansjoerg Kunc, Queen's University Belfast.

Philip Collins, (PhD student), funded by the University of Roehampton.
Do phenotypic traits explain reproductive skew in seabirds? (Field work undertaken on Puffin Island). Co-supervisor: Dr Jon Green, University of Liverpool.

Christle Coxon, (PhD student), funded by the University of Roehampton.
Overeating: The role of food composition, eating psychology and exercise on food intake. Primary supervisor: Dr Leigh Gibson, University of Roehampton.

Tessa van Walsum, (PhD student), self-funded (starting October 2014).
How can king penguins dive so deeply? A study of their cardio- and thermo-physiology. Co-supervisors: Dr Enrico Rezende, University of Roehampton; Dr Yves Handrich, CNRS Strasbourg.

Sarah Weldon, (PhD student), self-funded (starting October 2014).
Energy expenditure and the biopsychology of food perception during long-term caloric debt in ocean rowers. Primary supervisor: Dr Leigh Gibson, University of Roehampton.

Sinead Smith, (PhD student), funded by the Northern Ireland Department for Education and Learning (DEL) (starting October 2014).
Tracking the energetics of wild animals. Primary supervisor: Dr Michael Scantlebury, Queen's University Belfast; Co-supervisor: Dr Nikki Marks, Queen's University Belfast.

Christina Mulvenna, (PhD student), funded by the Northern Ireland Department of Agriculture and Rural Development (DARD) (starting October 2014).
Enhancing farm management by improving both commercial production and animal welfare: development and application of remote sensing technologies. Primary supervisor: Dr Michael Scantlebury, Queen's University Belfast; Co-supervisor: Dr Nikki Marks, Queen's University Belfast.

Membership of professional bodies

I am a co-opted member of the Animal Section of the Society for Experimental Biology

Teaching interests

I am programme convenor for the degree programmes Zoology and Biological Sciences.

I teach on the UG modules 'Physiology and Introductory Statistics', 'Biosciences Research Methods;, 'Animal Biology I', 'Animal Biology II' and the week-long 'Biosciences Field Course' based in Wales during late spring. I am also involved in teaching the Masters modules 'Ecological Methods and Practice', 'Research Methods in Biology', 'Health Research Methods' and 'Obesity: Risks and Prevention'.

I supervise a number of second year project proposals, third year independent studies, and third year projects, which often review and research issues of energy expenditure in humans, diving behaviour in animals and various non-invasive physiological topics.

Taught courses


Nutrition and Health


MSc/PGDip/PGCert Obesity: Risks and Prevention




Rey B, Halsey LG, Hetem RS, Fuller A, Mitchell D, Rouanet J-L (2015) Estimating resting metabolic rate by biologging core and subcutaneous temperature in a mammal. Comparative Biochemistry and Physiology Part A.


Reilly SM, Montuelle SJ, Schmidt A, Naylor E, Jorgensen ME, Halsey LG, Essner Jr RL (2015) Conquering the world in leaps and bounds: hopping locomotion in toads is actually bounding. Functional Ecology.

Halsey LG, Matthews PGD, Rezende EL, Chauvaud L, Robson AA (2015) The interactions between temperature and activity levels in driving metabolic rate: theory, with empirical validation from contrasting ectotherms. Oecologia. 10.1007/s00442-014-3190-5

Reeves S, Huber JW, Halsey LG, Villegas-Montes M, Elgumati J, Smith T (2014) A cross-over experiment to investigate possible mechanisms for lower BMIs in people who habitually eat breakfast. European Journal of Clinical Nutrition. 10.1038/ejcn.2014.269

Willener AST, Halsey LG, Strike S, Enstipp M, Georges J-Y, Handrich Y (2014) Reassessment of the cardio-respiratory stress response, using the king penguin as a model. Stress. 10.3109/10253890.2014.986451

Coward SRL, Halsey LG (2014) Energy expended during horizontal jumping: investigating the effects of surface compliance Biology Open. 3:815-820. 10.1242/​bio.20148672

Halsey LG, Tyler CJ, Kuliukas AV (2014) The energy cost of wading in water. Biology Open. 3:571-574. 0.1242/bio.20147831

Reeves S, Huber JW, Halsey LG, Horabady-Farahani Y, Ijadi M, Smith T (2014) Experimental manipulation of breakfast in normal and overweight/obese participants is associated with changes to nutrient intake and energy consumption patterns. Physiology and Behaviour.

Collins P, Green JA, Dodd S, Shaw P, Halsey LG (In Press) Predation of Black-legged Kittiwake chicks Rissa tridactyla by a Peregrine Falcon Falco peregrinus: Insights from time-lapse cameras. Wilson Journal of Ornithology. 126:158-161.

Lyons GL, Halsey LG, Pope EC, Eddington JD, Houghton JDR (2013) Energy expenditure during activity in the American lobster Homarus americanus: correlations with body acceleration. Comparative Biochemistry and Physiology A. 166: 278-284. 10.1016/j.cbpa.2013.06.024

Wilson RP, Griffiths IW, Legg PA, Friswell MI, Bidder OR, Halsey LG, Lambertucci A, Shepard ELC (2013) Turn costs change the value of animal search paths. Ecology Letters. 16: 1145-1150. 10.1111/ele.12149

Halsey LG (2013) The relationship between energy expenditure and speed during pedestrian locomotion in birds: a morphological basis for the elevated y-intercept? Comparative Biochemistry and Physiology A. 165:295-298. 10.1016/j.cbpa.2013.03.027

White CR, Cassey P, Schimpf NG, Halsey LG, Green JA, Portugal SJ (2013) Implantation reduces the negative effects of bio-logging devices on birds. Journal of Experimental Biology. 216: 537-542. 10.1242/jeb.076554

Halsey LG, White CR (2013) Comparative energetics of mammalian locomotion: humans are not different. Journal of Human Evolution. 63: 718-722.10.1016/j.jhevol.2012.07.008

Reeves S, Halsey LG, McMeel Y, Huber JW (2013) Breakfast habits, beliefs and measures of health and wellbeing in a nationally representative UK sample. Appetite. 60: 51-57.

Halsey LG, Huber JW, Hardwick JC (2012) Does alcohol consumption really affect symmetry perception? A three-armed placebo-controlled experimental study. Addiction. 107: 1273-1279. 10.1111/j.1360-0443.2012.03807.x

Knoss R, Halsey LG, Reeves S (2012) Ethnic dress, vitamin d intake, and calcaneal bone health in young women in the United Kingdom. Journal of Clinical Densitometry. 15: 250-254. 10.1016/j.jocd.2011.09.005

Robson AA, Chauvaud L, Wilson RP, Halsey LG (2012) Small actions, big costs: the behavioural-energetics of a commercially important invertebrate. Journal of the Royal Society Interface. 9: 186-1498. 10.1098/rsif.2011.0713

Halsey LG, Stroud MA (2012) 100 years since Scott reached the Pole: A century of learning about the physiological demands of Antarctica. Physiological Reviews. 92: 521-536. 10.1152/physrev.00031.2011

Qasem L, Cardew A, Wilson A, Griffiths I, Halsey LG, Shepard ELC, Gleiss AC, Wilson R (2012) Tri-axial dynamic acceleration as a proxy for animal energy expenditure; should we be summing values or calculating the vector? PLoS ONE. 7: e31187. 10.1371/journal.pone.0031187

Bidder OR, Soresina M, Shepard ELC, Halsey LG, Quintana F, Gomez-Laich A, Wilson RP (2012) The need for speed: testing acceleration for estimating animal travel rates in terrestrial dead-reckoning systems. Zoology. 115: 58-64 10.1016/j.zool.2011.09.003

Halsey LG, Watkins DAR, Duggan BM, (2012) The energy expenditure of stair climbing one step and two steps at a time: estimations from measures of heart rate. PLoS ONE. 7: e51213. 10.1371/journal.pone.0051213

Halsey LG, Stroud MA (2011) Could Scott have survived with today's physiological knowledge? Current Biology. 21: R457-R461.

Halsey LG, Jones TT, Jones DR, Liebsch N, Booth DT (2011) Measuring energy expenditure in sub-adult and hatchling sea turtles via accelerometry. PLoS ONE. 6: e22311.

Halsey LG, Shepard ELC, Wilson RP (2011) Assessing the development and application of the accelerometry technique for estimating energy expenditure. Comparative Biochemistry and Physiology Part A. 158: 305-314.

Lighton JRB, Halsey LG (2011) Flow-Through Respirometry applied to chamber systems: pros and cons, hints and tips. Comparative Biochemistry and Physiology Part A. 158: 265-275.

Halsey LG, White CR, Enstipp MR, Wilson RP, Butler PJ, Martin GR, Gremillet DR, Jones DR (2011) Assessing the validity of the accelerometry technique for estimating the energy expenditure of diving double-crested cormorants Phalacrocorax auritus. Physiological and Biochemical Zoology. 84: 230-237.

Halsey LG, Huber JW, Low T, Ibeawuchi C, Woodruff P & Reeves S (2011) Does breakfasting influence activity levels? An experiment into the effect of breakfast consumption on eating habits and energy expenditure. Public Health Nutrition. 15: 238-245.

Robson AA, Garcia de Leaniz C, Wilson RP, Halsey LG (2010) Behavioural adaptations of mussels to varying levels of food availability and predation risk. Journal of Molluscan Studies. 76: 348-353.

Hanuise N, Bost C-A, Huin W, Auber A, Halsey LG, Handrich Y (2010) Measuring foraging activity in a deep diving bird: comparing wiggles, oesophageal temperatures and beak-opening angles as proxies of feeding. Journal of Experimental Biology. 213: 3874-3880.

Halsey LG, Huber JW, Bufton RDJ, Little AC (2010) An explanation for enhanced perceptions of attractiveness after alcohol consumption. Alcohol. 44: 307-313.

Robson AA, Garcia de Leaniz C, Wilson RP, Halsey LG (2010) Effect of anthropogenic feeding regimes on the activity rhythms of laboratory mussels exposed to natural light. Hydobiologia. 655: 197-204.

Halsey LG, White CR (2010) Measuring energetics and behaviour using accelerometry in cane toads Bufo marinus. PLoS ONE. 5:e10170.

Halsey LG, Butler PJ, Fahlman A, Schmidt A, Bost C-A, Handrich Y (2010) Changes in the foraging behaviour and diving energetics of king penguins through summer and autumn: a month by month analysis. Marine Ecology Progress Series. 401: 279-289.


Halsey LG (2012) Narrow vision. Times Higher Education. (16th August).

Halsey LG (2012) Partying purges pounds. Complete Nutrition Focus. 4: 35-36.

Halsey LG (2011) The challenge of measuring energy expenditure: current field and laboratory methods. Comparative Biochemistry and Physiology Part A. 158: 247-251.

Reeves S, Huber J, Halsey L (2011) Breakfast and body mass index – why are breakfast eaters slimmer than breakfast skippers? Complete Nutrition Focus. 3: 9-11.

Halsey LG (2011) Optimal diving models: their development and critique requires accurate physiological understanding. Trends in Ecology and Evolution.

Reeves S, Halsey L (2010) Attaining energy balance with the must have toys this Christmas. British Medical Journal. 341(online). 10.1136/bmj.c7102

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