Thomas Henry Huxley Award and Marsh Prize

This Award is run in partnership with the Zoological Society of London and recognises a postgraduate research student whose thesis has made a significant contribution to a particular scientific field. Nominees are considered from a University of Great Britain and Northern Ireland, on the basis of original work, which has been awarded the degree of Doctor of Philosophy in the academic year in which the Award is presented.

The projects are judged by a panel of experts from the ZSL, who then select the winning study.

(2017 Awards were presented in 2018)

Pictured: A brown leaping frog in Kerala, India © ZSL

Dr Benjamin Van Doren 2021

Benjamin, University of Oxford, for his project ‘Flexibility in Avian Migration Across Scales’ 

This is a remarkable piece of work comprising six data chapters with a central theme of avian migration. This quality of this thesis is best summed up our Awards committee member: “Without exaggerating, I think this is the best thesis I have ever read (having been external and internal examiner for over 20 students and been the supervisor of another 25 that have successfully defended). There are no ‘filler’ chapters here, each represents a substantial contribution to the field. Indeed, I have already read many of them as they have been published (often in high profile journals such as Science, PNAS and Current Biology). Chapters 2, 4 & 5 in particular have had, and will continue to have, major impacts on the study of migration in birds (and migration in general) and Chapter 6 has recently been published and likewise well-received (I had also read this prior to receiving the thesis). The thesis combines existing data sets and field observations to produce a valuable contribution to our understanding of multiple, and often very different aspects of bird migration, all set within an evolutionary context. It is rare to receive a thesis of such quality, let alone one that covers deals with a topic in such a diverse and insightful way.

Previous Winners

Patrick Kennedy, University of Bristol


Patrick’s thesis considers two major, little-understood topics of biological relevance: how unpredictability in environmental conditions can impact the evolution of altruism (costly helping of others) and why some individuals in social wasp populations drift between nests (helping at colonies other than their own). The first of these questions is tackled using analytical modelling and evolutionary simulations, complex methods that 

Patrick learnt from scratch by seeking collaborations outside of his supervisory team. Answering the second question involved arduous fieldwork (in various Central and South American countries), experimental manipulations and sophisticated statistical analyses. The thesis contains several genuinely original ideas, and it excels in quantity and quality.  Patrick drove the direction of the research, suggesting innovative, insightful and ambitious ideas, devising carefully thought-out and often novel experimental and analytical plans to test them, and generating practical solutions to logistical challenges.

Erasmus Zu Ermgassen

Erasmus Zu Ermgassen, of the University of Cambridge, won this Award for his fascinating thesis – ‘Strategies for sustainable livestock production in Brazil and the European Community.’ Here Erasmus explores two means whereby livestock production could be made more globally competitive. The first is by re-legalising the use of food waste as pig food in the UK and the European Community. The second is by increasing the productivity of livestock systems in Brazil to reduce deforestation. Through his work, Erasmus has made a substantial and original contribution, and a large part of the work is already published in prestigious journals.

Shana Caro

Shana has won this Award for her thesis entitled ‘Social and environmental factors in the evolution of signalling’, focusing on signalling between parents and offspring.

This is a demanding and significant piece of scholarship which she approached with intellectual vigour and a positive attitude, and the reward is an extraordinary body of work.

Shana combines attention to detail and grasp of the “bigger picture”, which is rare in researchers at her career stage. She already has two papers in top international journals, one in Nature Communications and one in PNAS.

Shan Quah

Shan won this Award for her study entitled, ‘Conservation and innovation – the evolution of the metazoan microRNA landscape and its contribution to reproduction and development.’

Shan’s aim was to search for evolutionary ‘new’ microRNAs (miRNAs), to examine how miRNA genes originate and gain new functions relevant to animal evolution. She did extensive small RNA sequencing of all life cycle stages of two lepidopteran species. From the miRNA data that Shan generated, she worked out that there had been a ‘burst’ of miRNA invention in early lepidopteran evolution.

Shan took her paper further in two ways. Firstly, she discovers that one of the new miRNAs resides in the intron of an embryonic patterning gene and focused her attention on this, discovering a potential target gene for it in the same developmental pathway, which is a rare achievement. Secondly, she used the extensive data that she had gathered to formulate a revised model of miRNA evolution, which emphasises gene loss as much as gene gain.

All of the experimental and bioinformatics work in the paper was done by Shan and she provided the major intellectual input as well.

David Labonte

David won this Award for his PhD thesis entitled ‘Biomechanics of controllable attachment in insects’, undertaken at the University of Cambridge.

David’s thesis is extraordinarily informative and authoritative, and provides fundamental insight into the function of animal adhesive structures, combining new observations with clever experimentation and ingenious quantitative analyses of the underlying mechanisms. He collected extensive data from 225 species and his work has opened up new areas for research in this field.

David is a creative and independent scientist, who has come up with many new experimental ideas. During his PhD he has modified and improved complex systems, written his own software for most analyses and developed new mathematical models.

Dr Ben Ashby

Ben won this Award for his thesis on the role of genetic interactions and contact between hosts and parasites in a co-evolutionary context. Ben explores the diverse range of consequences of these roles, including the evolution of sex, genetic diversity and mating behaviour.

Ben’s thesis is entirely theoretical, but it is comprehensively informed throughout by biological questions and scenarios. His models are consistent with a variety of empirical data, and this is of crucial importance to evolutionary biology. The thesis is beautifully presented and four of the seven chapters are already published in excellent journals. There is no doubt that this study will have a great impact on the evolutionary-biology community.

Helen Leggett

Helen has been recognised for her thesis entitled: ‘Developments in social evolution and virulence in parasites.’, an exceptional piece of work which has generated some high-profile papers tackling major concepts in a novel and exciting way.  The thesis examines several different aspects of host-parasite interactions through five main chapters.  The other two main chapters include a comparative study of the relationship between immune system subversion and pathogen virulence, and an experimental study looking at the evolution of virulence in a spatially distributed population.

Helen demonstrated that, contrary to standard theory on virulence evolution, pathogens with higher virulence do not tend to have higher within-host growth rates. Given the importance of this assumption in standard theory on virulence evolution, this finding is likely to be of great significant to researchers in this area, and it has the potential to change the way people think about the subject.

Mary Caswell Stoddard

Mary’s thesis dives into the evolution of avian egg features, mainly colour and pattern, to test various hypotheses for their observed features. She used the full range of colour vision of birds, which have UV detection in order to understand colour and pattern. Her study has shown evidence that species that host cuckoos have experienced selection to make their eggs more recognisable, which has proved to be a novel finding.

Mary has deployed a great diversity of techniques in her thesis. Her work on egg colour and pattern uses detection approaches and numerical methods derived for avian vision, some of which have been invented by her. Her work testing the sexually-selected eggshell colouration theory involved a great-tit field season with a double-cross fostering design, while her investigations into whether pigment strengthens eggshells against injury involved work in a mechanics lab. Mary has 11 publications in Web of Knowledge, including several of the chapters of her thesis.

Kate Jordan

Kate, who studies at the University of Warwick, won the Award for her thesis entitled ‘Exploring the generative architecture of intramembranous ossification’. Her experimental work in developmental biology has revolutionised thinking about vertebrate skeletal development and hard-tissue evolution.

Kate used a variety of techniques in her study, including 3-D reconstruction, to provide conclusive evidence that the current model of how the bones of the skull is entirely wrong. Her research impacts a number of fields including veterinarians, human surgeons, bioengineers and evolutionary biology.

Sam Weber

Sam is a student at Exeter University and has been recognised for his study of Maternal effects in the green turtle. He explored two questions of fundamental and applied significance – is variation in the levels of maternally-derived antioxidants in eggs an adaptive response by mothers to reduce oxidative stress in their offspring? And is natal homing behaviour adaptive?

Sam focused on the green turtles of Ascension Island and used a range of observational, experimental, biochemical and statistical techniques to derive answers to both questions. His work makes a novel contribution to knowledge on the factors which influence variation in egg quality and composition in turtles and the consequences for the hatchlings.

Tom Fayle

Tom has been selected for this Award for his thesis ‘Ant community structure in a rainforest microcosm’, which is based on an empirical and statistical investigation of the biodiversity and community structure of ants living in tropical rainforest ferns. Tom combines excellent natural history with experiments and computer simulations to determine the rules that might determine species assemblies. His results are clear and well presented and the study also reflects the fact that Tom managed to learn the means of identifying 71 different species of ant, a notable achievement in itself.

Hannah Rowland

Hannah was recognised for her exceptional thesis titled ‘The visual and behavioural ecology of countershading and other prey defences’. Her work has already resulted in research papers to be published in Nature, Philosophical Transactions of the Royal Society, Proceedings of the Royal Society and Animal Behaviour.

In her study, Hannah explores visual deception in prey to reduce the risk and incidence of predation, focusing mainly on crypsis and concluding with a very neat study of mimicry. Her experimental studies use avian predators and prey ranging from live Lepidoptera larvae to artificial food, and are carried out in environments ranging from semi-natural to aviaries.

In her work, Hannah convincingly resolves a long-running debate and shows that the net effect is always mutualistic, not parasitic, for prey with unequal defences once prey densities are incorporated. She also demonstrates that, in Batesian mimicry, the existence of a palatable mimic does not necessarily diminish the net survival of a toxic model ‘prey’ population, even when the mimic is at high densities.

Tim Hawes

Tim explored the processes that enable certain arthropods to inhabit the exceedingly cold ecosystems of terrestrial Antarctica. Tim’s research forms a logical sequence of nine complementary studies, most combining laboratory and field experiments. They range from pure physiology to ecology, and are distinguished by the originality and appropriateness of each hypothesis being tested and by the neatness and rigour of the experimental procedures and analysis. Most make real advances in the understanding of a process rather than consolidate knowledge through repeating old experiments on new species.