Insectes Sociaux Best Paper 2019

It is a pleasure to announce that the winner of the 2019 Best Paper Award is “Vitellogenin and vitellogenin-like gene expression patterns in relation to caste and task in the ant Formica fusca” by Claire Morandin, Anna Hietala and Heikki Helanterä at the University of Helsinki, Finland. Here is the journey of how this project came to be, in Claire’s words.

When I started my Ph.D. in early 2011, Vitellogenin (Vg) was the “hit” gene that everyone was interested in. Vg is one of the most studied genes involved in the division of labour across social insects. My first Ph.D. project was to look at Vg expression patterns across multiple Formica ant species, but upon constructing the first Formica transcriptome (Dhaygude et al., 2017), we surprisingly came across not only one Vg gene, but four. To understand these new Vg-like-genes (as we named them), we performed gene expression analyses, protein modelling, and evolutionary analyses, and found that these three new homologues were the result of ancient duplications. They partly differ in their conserved protein domains and have undergone rapid evolution after duplications. Furthermore, their expression patterns and thus their likely roles in social regulation were not consistent across the seven Formica species we looked at, providing important new insights into the complexity of insect social behaviour and gene expression variation amongst even closely related species.

unnamed (3)

Understandably, we got really excited about these Vg-like genes and knew we needed to look deeper into their roles. In species with multiple conventional Vg (such as Solenopsis invicta), the multiple copies show sub-caste and task-related expression patterns, potentially linked to the loss of reproductive constraints and evolution of new functions for the duplicated copies (Wurm et al., 2011). We then questioned if similar caste- and/or task-related expression differences would likewise have emerged during the ancient duplication events between/among the conventional Vg and the three Vg-like genes. Hence, to begin to comprehend the role of Vg homologs and their potential involvement in division of labour, we designed a study that would allow us to investigate the relationships of expression patterns not only between castes (queens vs. workers), but also between different tasks (nurses vs. foragers), between colonies with and without a queen, and between several points in time.

At the same time, Anna started her master thesis with us. It was a challenging and stimulating project that involved a lot of field and lab work. Once the snow melted, we went out around Tvärminne Zoological Station in the south of Finland and collected dozens of F. fusca colonies. The time frame was tight, as we needed to find them after they came out of hibernation, but before they got too active, in order to catch the entire colony (or most of it), the queens included. As we needed colonies with several queens, we could not afford to miss queens while collecting. Back in the lab, the colonies were carefully sorted to find and count all of the queens (that’s a tedious job that involves going through a bucketful or two of nest material and soil for each nest, and hundreds or even a few thousand workers that do not appreciate your efforts). Experimental nests were established in plastic trays with a feeding platform so we could differentiate nurses and foragers. The experiment lasted for 20 days; every five days we collected nurses, queens, and foragers from each nest for gene expression analyses and checked whether queens or workers had been laying eggs. After that, we brought back the samples to the University of Helsinki for gene expression analysis. Anna extracted RNA from more than 500 individuals and performed qPCR analysis (needless to say it was a challenging task for someone who has never done any lab work after one or two basic courses, but she managed brilliantly). Apart from the wet lab work, Anna also dissected the ovaries of a few hundred workers to see whether queenlessness incites ovary development in workers.

Our results showed that each of these genes had a unique caste-specific expression pattern in F. fusca. Expectedly, we found a significant caste and worker task-related increase for the conventional Vg. We found that task (nurses vs. foragers) was the only factor that explained expression variation among workers in any of the studied genes and that removing the queens did affect expression, despite the fact that the proportion of fertile nurses increased significantly. As in previous studies (Kohlmeier et al., 2018; Salmela et al., 2016), our results are consistent with the idea that Vg-like-A may be involved in worker behaviour, and Vg-like-B in stress resistance in ants, while Vg-like-C displayed a consistent forager-biased expression pattern (just like in other social insects (Harrison et al., 2015), suggesting that Vg-like-C might have sub functionalized to a completely different role.

With this project, we aimed to get a clearer picture of the roles of these newly found Vg-like genes for caste differentiation. We still do not know their precise roles, and for example, tissue-specific expression analyses would be an important next step, but at least now we know that their expression patterns are consistent with roles in the division of labour separate from the conventional Vg. We hope this study will spark further interests in these really interesting Vg homologues, and hopefully, at some point we will find out exactly what these genes are doing at a molecular level.

As many projects do, this one took its time as well, and there were challenges along the way. For example, extra effort was needed from Anna who had to write the thesis in English and not in her native Finnish as Claire as her supervisor needed to understand it as well. It was a long journey from the beginning to Anna’s MSc thesis, and then publication in Insectes Sociaux. After all the work for this study, the award means a lot to all of us!


Dhaygude, K., Trontti, K., Paviala, J., Morandin, C., Wheat, C., Sundström, L., & Helanterä, H. (2017). Transcriptome sequencing reveals high isoform diversity in the ant Formica exsecta. PeerJ, 2017(11), 1–31.

Harrison, M. C., Hammond, R. L., & Mallon, E. B. (2015). Reproductive workers show queenlike gene expression in an intermediately eusocial insect, the buff-tailed bumble bee Bombus terrestris. Molecular Ecology, 24(24), 3043–3063.

Kohlmeier, P., Feldmeyer, B., & Foitzik, S. (2018). Vitellogenin-like A–associated shifts in social cue responsiveness regulate behavioral task specialization in an ant. PLoS Biology, 16(6), 1–26.

Salmela, H., Stark, T., Stucki, D., Fuchs, S., Freitak, D., Dey, A., … Sundström, L. (2016). Ancient duplications have led to functional divergence of vitellogenin-like genes potentially involved in inflammation and oxidative stress in honey bees. Genome Biology and Evolution, 8(3), 495–506.

Wurm, Y., Wang, J., Riba-Grognuz, O., Corona, M., Nygaard, S., Hunt, B. G., … Keller, L. (2011). The genome of the fire ant Solenopsis invicta. Proceedings of the National Academy of Sciences of the United States of America, 108(14), 5679–5684.

We’re seeking a new Social Media Editor

I want to take this opportunity to express my heartfelt thanks to all of the blog contributors and interviewees for providing some brilliant blog content. The past year has been big for the blog, with over 7,000 readers (in 2019) from all over the globe. Our other Social Media Editor, Bernie Wittwer, is leaving her position. We will be seeking her replacement in the next few months.

Part of what makes working for Insectes Sociaux special is that it is a truly international journal, and the success of the journal depends on global science conducted by individuals at all career stages, all over the world. As the Social Media Editor, I seek contributors to the blog that are as diverse as the contributors to the journal. These contributors not only increase the representation of all groups in the diverse social insect community but also increase the impact of the blog, as more social media users share the blog with the members of their increasingly global networks. I am proud to be a part of this and work to continue this diversity in the future.

For those of you who may be interested in becoming the next Insectes Sociaux Social Media Editor, I offer a brief description of how the role works for me. For about two hours a week, I spend my time contacting potential blog contributors and social insect scientist interviewees, managing social media (note – our next Social Media Editor will need to have a Facebook account), finding images and videos to complement the blog posts, and laying out the blogs for publication on the WordPress site. But most of all, I spend time editing the blog posts, working with the authors to present their research and their experience doing it in the clearest and most engaging way for a non-expert audience. I aim to make the science accessible and to help the blog contributors find their voice.

If this sounds like something that you might want to take on, please get in touch with me at or Dr. Miriam Richards, Insectes Sociaux Editor-in-Chief, at to express your interest.

Thanks again everyone,


Interview with a social insect scientist: Robert J. Warren II


IS: Who are you, and what do you do?

RW: My interest is in the impact of global change (species invasion, climate change, and habitat fragmentation) on species interactions, which quite often steers me toward social insects.

IS: How did you develop an interest in your research?

RW: Really, at its core, my interest in research stems from walks in the woods. My favorite research projects tackle natural history observations with ecological theory.

IS: What is your favorite social insect, and why?

RW: The Aphaenogaster rudis complex (woodland ants in eastern U.S. deciduous forests) certainly are my favorite social insects because I spent much of my life hiking and playing in woodlands, and I never knew that a single species was so dominant (both in abundance and impact). Now, I see them everywhere.

IS: What is the best moment/discovery in your research so far? What made it so memorable?

RW: I spent my doctoral research trying to explain the contrasting distribution of two myrmecochorous plants in the Southeastern U.S. based on the niche requirements of the plants. One day, when loading equipment into the back of my truck, I thought, ‘what if it is the ants?’ My research had solely focused on plants up until that time, and incorporating ants opened up a whole new world for me (including new insights into global change ecology).

IS: Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

RW: I teach ecology and biostatistics in a biology department (SUNY Buffalo State). I share my research and field experience with students – most of whom want to go into the medical field and do not think ecology is relevant to their careers – to connect with the students and help them see a different aspect of biology.

IS: What do you think are some of the important current questions in social insect research, and what’s essential for future research?

RW: I am fascinated with how often social insects get cheated. We know of many great benefits that come with sociality/eusociality, but one of the main benefits seems to be that a colony can amortize the cost of being cheated across many individuals and hence tolerate it more easily than a solitary organism.

IS: What research questions generate the biggest debate in social insect research at the moment?

RW: In my little area, ant-mediated seed dispersal has long been described as a mutualism despite little evidence supporting palpable benefits for the ants. I have received surprisingly strong and emotional negative responses, particularly from plant-oriented folks, when questioning it as a mutualism.

IS: What is the last book you read? Would you recommend it? Why or why not?

RW: Last Train to Memphis, by Peter Guralnick. I would recommend it. Guralnick does an amazing job detailing the rise of Elvis Presley, including showing that Presley had a deep understanding and passionate love for black music, and you realize that a lot of the resistance to Presley in his early years was racist resistance to his integrating black musical styles into what became rock and roll.

IS: Outside of science, what are your favorite activities, hobbies, or sports?

RW: I love hiking and all that stuff, and I spend a lot of time doing home improvement on our 19th Century Victorian home. I also love watching Indiana University basketball and University of Georgia football.

IS: How do you keep going when things get tough?

RW: I have made a few great decisions in my life, and my wife of 27 years is one. She is my foundation.

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

RW: These days that question is a little open given that we can bring a Kindle and mp3 player with thousands of options, but I will try and answer in the traditional sense. I would bring Alice in Chains “Grind” because I can listen to it again and again, Hermann Hesse “Siddhartha” because I seem to find something new every time I read it, and a large supply of Tabasco sauce because I like it on my seafood.

IS: Who do you think has had the most considerable influence on your science career?

RW: Retired University of Georgia Professor Ron Pulliam. I entered his lab as a former newspaper journalist and left a scientist.

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

RW: Work with people that you like and find projects for which you have passion because it is a helluva lot of work to do for low pay and little reinforcement.

IS: What is your favorite place science has taken you?

RW: As a literal ‘place,’ I love doing work in the Southern Appalachian Mountains where, even though there are thorns, wasps, and steep slopes, I can work in shorts and a t-shirt and get lost in the green. As a figurative ‘place,’ I love working with students and scientists, which is endlessly fascinating and rewarding.

Interview with a social insect scientist: Tim DeLory


IS:Who are you, and what do you do?

TD: My name is Tim DeLory. I am currently a Ph.D. student at Utah State University in Dr. Karen Kapheim’s lab, where we study the evolution of sociality in bees. The objective of my dissertation is to study genome evolution in bees across a range of bee social life histories using a bioinformatic approach. I am particularly interested in the potential impact that these complex social phenotypes can have on the evolution of genomes.

IS:How did you develop an interest in your research?

TD: I first learned about social insects in my undergraduate entomology class. My professor, Kevin Alexander, explained the haplodiploid mating system of hymenopterans to us and showed us some of the different species of bees with variable sociality. This piqued my interest in social insects. The following summer, I had the opportunity to attend a Research Experience for Undergraduates (REU) at UNC Greensboro under the guidance of Olav Rueppell, where we studied recombination rates in honeybees. It was after this REU that I knew I wanted to make studying social insects my career path and use bioinformatics approaches to do this.

IS:What is your favorite social insect, and why?

TD: I would have to say Meliponini. Maybe it’s cheating to say a whole tribe is my favorite, but it’s their diversity of behavior and life histories that fascinates me. They have a wide variety of castes and nesting strategies. I have also always loved reading about interspecific interactions between eusocial species, and the ​Lestrimelitta limaois a fascinating example of this. They have a “robber” caste instead of a foraging caste which robs the reserves of other stingless beehives. The pollen-collecting morphology of ​Lestrimelitta is reduced and vestigial. It is interesting to me that the forager caste is not inextricably linked to pollen-collecting in bees. On a personal note, I think they are beautiful (especially ​Tetragonisca angustula) and have an incredible variety of kinds of honey.

IS:What is the best moment/discovery in your research so far? What made it so memorable?

TD: I would say the most memorable moment in my research was when I was able to see the results of a simulation that I made when finishing a fire ecology project I had been working on before starting my Ph.D. It relied on some field data we had collected as well as GIS data that other students had gathered. The results were interesting, but the moment was memorable because I liked the collaborative effort that went into it. I was able to really integrate my math background, as well. It gave me a strong sense of affirmation that research in the life sciences was the right career path for me.

IS:Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

TD: I am a teaching assistant for some introductory biology labs at my university. I try to discuss my research whenever appropriate in these classes to demonstrate that people do apply the skills they are learning. When teaching, I also try to expose my students to other career opportunities beyond academia or medical school, such as land management agencies or non-profit work. I have some friends in these areas that I can refer them to if they have questions. I think this helps students feel less siloed by a biology degree, which hopefully translates to retention. As far as conversations with the community at large, I really consider it a victory when I can explain that there is a lot of research beyond honeybee health going on in entomology, and why basic scientific research, in general, is valuable.

IS: What do you think are some of the important current questions in social insect research, and what’s essential for future research?

TD: I am relatively new to research in social insects and research in general. But I would say some of the recurrent issues I have noticed have to do with the rate of advancement of sequenced genomes, while the characterization of insect life histories has not advanced nearly as rapidly in recent years. In 06’, the honeybee genome was assembled, and there was a lot of information about its social life history to contextualize that genome. We are now in a position where we can afford to sequence bees whose more basic qualities are still a mystery. It will be interesting to see how new assessments of bee life histories can help to direct sequencing efforts. Or perhaps as sequencing becomes more and more affordable, we will sequence results to direct which bees we select to do behavioral studies in the field.

IS: What research questions generate the biggest debate in social insect research at the moment?

TD: I know that the value of kin selection theory as a framework for understanding social evolution has been repeatedly brought into question over the years, and has sparked many debates. Although I am not sure if this issue is as contentious today. One recent question that seems to generate a lot of discussion and differing opinions is the role of using insects with semi-social or facultatively eusocial lifestyles to understand eusocial evolution as a whole. I think the research community has not yet reached a consensus on what we can reasonably infer about the evolution of extant eusocial lineages using less socially complex insects as ancestral proxies.

IS:What is the last book you read? Would you recommend it? Why or why not?

TD: The last book that I read was called The Dragon Reborn. It was a fun read for me. I would recommend it if you really like reading Tolkein-like fantasy books with expansive world building and have read the first two books of the Wheel of Time series. I would not recommend it if you are interested in a book with deep or dynamic characters. The plot moves pretty slowly as well because it is part of a much larger series.

IS:Outside of science, what are your favorite activities, hobbies, or sports?

TD: I like different outdoor activities such as hiking, rock-climbing, and skiing. I also enjoy jiu-jitsu.

IS:How do you keep going when things get tough?

TD: Mostly, I just think about what got me interested in social insects in the first place. The idea of these various insect societies interacting and inhabiting the planet and acting out these amazing and intricate narratives reminds me of a science fiction novel. It’s endlessly fascinating. It’s easier to push through when I can remind myself that my job is to research something that is fundamentally cool.

IS:If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

TD: I would bring seeds. I used to have a large vegetable plot when I was younger, and I loved tending to it. So, I think gardening would be fun, and an excellent way to have a steady source of food. I would bring my collection of books. I think I could read Frank Herbert’s Dune once a year and not get sick of it. Lastly, I would bring a volleyball, because it is always good to have a friend.

IS:Who do you think has had the most considerable influence on your science career?

TD: I had a fantastic group of professors during my undergrad at Western Colorado University (WCU) who all encouraged me to do research and taught me valuable skills to do my research. My research advisor at WCU, Dr. Jonathan Coop, really comes to mind. He gave me a lot of creative control over designing a simulation for a project of his that I was working on. I felt like a colleague that was making a real contribution. Beyond that, he has always respected my personal decisions, including leaving school to work when I needed a break.

IS:What advice would you give to someone hoping to be a social insect researcher in the future?

TD: I would say that reading a healthy amount of papers is helpful for this in the beginning. The literature will give you a sense of what intrigues you about social insects. You can check out the lab sites of the authors of the papers that you especially liked, and contact those authors. That is how I met my Ph.D. advisor. It has been my experience that everyone I have contacted in the social insect community is affable. They will recommend other labs, too, if they think those labs could be a good fit for you.

IS: What is your favorite place science has taken you?

TD: I had the opportunity to go to an old wildfire site from the 2002 Rodeo-Chedeski burn in Arizona to collect data for a fire ecology research project. I was on top of a mesa looking around, and it really made me realize how expansive the burn was. That vista definitely made a strong emotional impression. I could also see how the landscape had such a dynamic recovery response from one drainage to the next during the time I spent there.

Interview with a social insect scientist: Antoine Felden

IS: Who are you and what do you do?

AF: Hi, I’m Antoine Felden, and I live in Wellington, New Zealand. Currently, I study the Argentine ant immune system in response to pathogens, mostly using transcriptomics and functional genetics (RNA interference). I also have other ongoing projects, too, like the study that just published in Insectes Sociaux about how a mite might be associated with wasp pathogens. I’m getting more and more experience in computational biology, and that is what drives me. I love fieldwork (and bench work to some extent, too) but I thrive working with computers and digging deep into datasets.


Probably talking about Toulouse and bees at the Biology & Genomics of Social Insects conference © Cold Spring Harbor

IS: How did you develop an interest in your research?

AF: I have had a passion for ants as far back as I remember. That is what got me into biology in the first place. I worked on a lot of different topics over the years, from behaviour to pathogens and immune response, locomotion mechanics, learning, and nutrition, all using ants as study models, and I’ve always been fascinated by what I was doing. So I suppose the short answer to that question is: ants.

IS: What is your favorite social insect and why?

AF: I suppose I don’t need to say it twice! I think ants tap into the fascination I have for miniature worlds. I was a pretty weird kid staging up small sceneries and playing with little things until an unusually old age, and ants perfectly fit this kind of aesthetic. Then, growing up and studying, I learned about the complexity of their biology, and a more scientific fascination about them emerged. I am amazed by all the different characteristics that have evolved in different species, from kidnapper ants to fungus-growing ants, army ants, aphid-tending ants, unicolonial ants, and so one. There is so much diversity in the ant world that one can never stop being amazed.

More anecdotally, there was a colony of Formica selysi living in the sandy backyard of the family holiday house on the Atlantic coast of France. After looking forward to seeing them again every year in summer, I have to say they have a special place in my heart; they looked stunning with their iridescent silver cuticle, running in the sun.

IS: What is the best moment/discovery in your research so far? What made it so memorable?

AF: One highlight discovery was when I was to study appetitive conditioning in Argentine ants to decipher nestmate recognition mechanisms, back in Berkeley with Ellen van Wilgenburg and Neil Tsutsui. I was using something similar to the bee “proboscis extension reflex” in ants to figure out if there were able to learn some olfactory cues better than others. Watching them under the stereomicroscope sticking their tongue out as they remembered that a given smell was followed by a sweet treat was a successful experiment like few happen. That was a while ago; now, my academic path is taking me to fuzzier transcriptomic studies where discoveries often bring more questions than answers.

IS: What do you think are some of the important current questions in social insect research and what’s essential for future research?

AF: Questions that revolve around caste differentiation and its molecular basis have the potential to have significant implications for our understanding of developmental biology as well as the evolution of sociality. For our society’s benefit, there might also be a lot to learn in studying aging in ants where queens can have such an extended lifespan compared to the average worker. What is essential for further research in most fields is to develop more models for functional genetics, like in the CRISPR-ant studies published in the last couple of years. It will be paramount to investigate these processes better, and I’m really excited about this!

I also think that unicoloniality that some ant species display is an odd phenomenon and that its ultimate mechanisms are still poorly understood. What caused the formation of these large supercolonies of ants? What are the behavioural consequences of such an upscale colony size? Is unicolonial social structure viable over evolutionary time? I think these questions still need more attention.

IS: What research questions generate the biggest debate in social insect research at the moment?

AF: I see that there is still a lot of room for debate in themes revolving around the evolution of sociality. The recent papers by Boomsma & Gawne (2018) on superorganismality and Linksvayer & Johnson (2019) about misconceptions on social complexity were pretty thought-provoking and highlight how much there is still to understand.

IS: What is the last book you read? Would you recommend it? Why or why not?

AF: First, the series (especially the first volume) called Les fourmis by Bernard Werber is a must-read for science fiction enthusiasts and naturalists alike. It is a very well-read novel, and there are still some accurate science facts in there (if one lets scientific rigour loose a little). I also love most of Virginie Despentes’ work. She writes about lives going on destructive train wrecks. Her characters are often objectively dark and despicable people, but following their struggles from their own perspective makes you uncomfortably sympathetic with their twisted journeys. I definitely recommend checking them out; there are English translations available!

IS: Outside of science, what are your favorite activities, hobbies or sports?

AF: Since I moved to New Zealand, I took up surfing more seriously. I’m lucky to live in Wellington; there is a nice beach easily-accessible by bus (I’m one of these university people who don’t have a driving licence… I know I’m not the only one out there!). It’s often a mix of chance and negotiation skills to hop on the bus with a surfboard, but it’s always worth a try. And thanks to the Argentine ants that like it hot and sunny, fieldwork in the warmer northern tip of New Zealand is conveniently located around beautiful surf beaches. Beyond that, I really like hanging out with my friends.


Collecting Argentine ants living inside beehives in the Far North of New Zealand, with Jana Dobelmann © Phil Lester

IS: How do you keep going when things get tough?

AF: Highs come with lows, and I try to remember that problems eventually drift away just like clouds in the sky. When issues arise, I try to not lose sight of their resolutions that come with trying a bit harder.

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

AF: I’d bring a sound system, a laptop, and solar panels – provided that supplies are already on the island. Then I’m all set to take science notes, write a book, then celebrate the outcomes with a dance.

IS: Who do you think has had the most considerable influence on your science career?

AF: My first internship mentor – Tim Brütsch, in Lausanne, who was doing his Ph.D. at the time. He taught me the necessary dedication for experimental work, how the academic world works, and sparked my interest in an evolutionary perspective on biology. I’d also like to mention my dear colleague Raphael Aguillon (not a social insect researcher – but no one is perfect) who asks the big questions and also showed me what passion for science looks like.

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

AF: Stay around good people and trust that with time, luck always strikes. I have both of those at Victoria University working with Phil Lester: a pleasant working environment with good funding to answer our questions, as well as having been at the right place and the right time.

IS: What is your favorite place science has taken you?

AF: I went to beautiful Tokelau to deal with a Yellow Crazy ant invasion as part of Monica Gruber’s Pacific Biosecurity program. Tokelau is a tiny atoll nation north of Samoa. It’s pretty hard to get to, as it is only accessible by boat every fortnight from Apia, and it is one of the most incredible places I’ve seen, full of lovely people. To this day, I can still reminisce how constantly stoked I was to be there.


Telling Tokelau kids about invasive ants © Evan Brenton-Rule

Book review: “660 Photographs of Termites and Their Control” by David Mora del Pozo

Review by Thomas Chouvenc


When you see someone passionate about a topic, it can humble you. Especially if this topic happens to be the one that you are also passionate about. Let me explain.

Recently, 660 Photographs of Termites and Their Control, by David Mora del Poso, was released, and I was fortunate enough to receive a physical copy of it. As I opened the book and gently flipped through the pages, my mind was immediately blown away by its beauty, its inherent poesy, and its deep dive into termite biology and complexity.


One thing that will be quickly apparent to you is that this book is not your usual lengthy textbook about the biology of a given bug. It is instead a fascinating compilation of photos that David took over decades during his time addressing termite pest problems around the world.  Each picture published in this book tells us a fantastic story about the biology of termites, their inherent beauty, and their abilities. However, the storytelling is not performed through a dry academic approach; instead, it immerses you within live termite colonies using remarkable macro shots. It also opens your eyes to the rather difficult relationships that termite societies and human societies sometimes have.


For a 425-page book that weighs a little bit more than 2 kg, it sure is a heavyweight. However, the 400 first pages are a compilation of (amazing) pictures with reference numbers. On the last 25 pages, you will find a text corresponding to each of the 660 images found in the first 400 pages providing an explanation of what you see. The text is provided both in English and Spanish, which is a remarkable effort from David to reach a wider audience, as his proficiency in English is limited.

As you open the book randomly, you will be glued to the first page you see, then flip back and forth, until you realize you just spent 15 minutes doing so. This book does this to you. In the process, you are immersed in the biology of various termite species, primarily with infamous pest species within Coptotermes, Reticulitermes, Cryptotermes, Nasutitermes, Heterotermes, and a few other genera that David encountered during his work.

Screen Shot 2019-11-15 at 2.37.41 PM

Macro shots will take you through the journey of the molting process of a worker, the complexity of the hatching process from an egg, the wing bud development and sclerotization of an alate, the transformation of a pre-soldier. Wider shots will show you the incredible complexity of nest structures, the intensity of swarming events, and the potential damage termites can do to structures and precious historical human artifacts.

Screen Shot 2019-11-15 at 2.38.35 PM

This book also provides a unique insight into the world of termite control – not always the most visible or respected in the world of academia. With $40+ billion in yearly damage worldwide, termites are usually unaccepted in someone’s house, which is an attitude you cannot ignore as a termite researcher. The impact of termites on people is beautifully portrayed in David’s book. It reflects what can happen if some of the termite pest species interact with human societies.

David has spent more than 25 years dealing with termite pest problems around the world, and he has been instrumental in accomplishing something that no one else has accomplished. Since the advent of Chitin Synthesis Inhibitor Baits for subterranean termite pest species in 1995, there has been a fundamental change in the philosophy for termite control around the world. Instead of spraying a large amount of pesticides around structures and hoping for the best, baits now aim at eliminating the whole colony after termites feed on a ridiculously small amount of a termite-specific formulated baits. Beyond this cultural shift, area-wide baiting programs were proposed to create large termite-free zones. Such programs have been implemented in various places around the world, with mixed results, primarily because of logistical difficulties that are inherent to cultural hurdles of human political landscapes. The problem is often not the termite pest – it is how people are handling them and how conflicts of interest may defeat the attempt.

Screen Shot 2019-11-15 at 2.39.50 PM

David is an exception. He successfully implemented a city-wide termite control program in several cities across Spain. Some of these cities have not seen termite damage since their implementation. This is quite remarkable, but what is even more impressive is that David documented the extensive implementation of these programs and provided a large chunk of these data in his book. Yes, there is a section that is dedicated to how he implemented area-wide termite control. To be honest, the work he documented leaves an amazing roadmap of how termite control should be done, with the utmost respect for the environment.

Screen Shot 2019-11-15 at 2.40.47 PM

David Mora Del Pozo may not be well known from social insect research groups. Yet, with this book, David is leaving a unique legacy to inspire many future budding entomologists. On top of this, 660 Photographs of Termites and their Control has all the attributes you need for a proper display on a coffee table or as an entomo-gift for the holidays.


More about 660 Photographs of Termites and their Control at

Interview with a social insect scientist: Giselle Martins Lourenço

IS: Who are you and what do you do?

GL: My name is Giselle M Lourenço, and I recently finished my Ph.D. in Ecology at the University of Campinas (Brazil). Currently, I am a postdoctoral researcher at the Centre for Ecological Synthesis and Conservation at the University of Minas Gerais (Brazil). My research aims to understand the distribution and movement of insects in space and time and how environmental filters and climate change influence them. During my master’s degree in ecology at the University of Ouro Preto (Brazil), I studied the distribution of herbivorous insects and ants in the canopy. It was at this moment that the opportunity arose to develop, with my dear friend Fabíola Keesen (Ph.D. student at the University of Rio de Janeiro, Brazil), an experiment to evaluate the recruitment of the ant Dorymyrmex thoracicus and a computer model for the trail formation. We reported this study in a recent paper at Insectes SociauxRecruitment and entropy decrease during trail formation by foraging ants (Lourenço GM, Keesen F, Fagundes R, Luna P, Silva AC, Ribeiro SP, Arashiro E, 2019).


Me and my friend Fabiola Keesen conducting an experiment with the ant species Dorymyrmex thoracicus (Dolichoderinae) (Brazilian Atlantic Forest). Image: Reisla Oliveira

IS: How did you develop an interest in your research?

GL: I developed an interest in ecology and insects during my undergraduate degree, but I started studying ants during my master’s. My friend Fabíola Keesen studied biological modeling, so we decided to integrate our knowledge to work together. The idea of our recent paper at Insectes Sociaux was to integrate our field experiments with computer modeling, which allowed us to extrapolate and understand ant foraging. Further, we invited our advisors (Dr. Sérvio P Ribeiro and Dr. Everaldo Arashiro, respectively) and other expert collaborators on these studies (Roberth Fagundes, Pedro Luna and Alcides C Silva). It was very productive to work in an interdisciplinary way, allowing new experiences for all co-authors.


Ant species Dorymyrmex thoracicus Gallardo (1916) as a study organism for investigating the recruitment and entropy decrease during trail formation (our recent paper at Insectes Sociaux). Image: Reisla Oliveira

IS: What is your favorite social insect and why?

GL: Ants are my favorite social insect. It is incredible that they can organize themselves to perform different activities throughout the day, balancing the costs and benefits for the entire colony with each decision.

IS: What is the best moment/discovery in your research so far? What made it so memorable?

GL: My Ph.D. defense! It was a fantastic moment, a closure of four years very well lived! Among all the challenges I experienced during the doctorate, I had the opportunity to learn a lot, make great friends, and visit amazing places. Finishing this step was hard because I did not want it to end! However, it was great to share this achievement with friends and my lovely family!

IS: Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

GL: Whenever I talk about insects and their behavior, especially ants, I try to show how small organisms can be very organized and ready to act in different situations. The examples associated with foraging, colony defense, and recruitment strategies arouse curiosities in everyone, from biology students to non-specialists in the field. I try to participate in events for the general public, bringing curiosities about insects and their role in our daily lives.

IS: What do you think are some of the important current questions in social insect research and what’s essential for future research?

GL: I would think that it is important to connect field experiments with computer modeling. These methods can help to demonstrate that the organization of complex systems is also a quality of ant colonies. Employing this complex systems approach, we can explain collective behavior that cannot be detected by studying individual and isolated ants.

IS: What research questions generate the biggest debate in social insect research at the moment?

GL: Questions related to the effect of climate change on social insect behavior. Also, the discussion about the threat to insect ecosystem services due to intensive pesticide use has been growing and drawing the attention of the general population.

IS: What is the last book you read? Would you recommend it? Why or why not?

GL: Lab Girl, by Hope Jahren. I got this book from some friends after my doctorate defense and was a very special read for the moment I am currently living. Hope narrates her story from the beginning of her career and talks a lot about the challenges of young PhDs who seek the dream of growing up and establishing themselves as a teacher and researcher. I highly recommend it, especially for young women scientists!

IS: Outside of science, what are your favorite activities, hobbies or sports?

GL: I like to dance the samba, travel, be with my family, and practice yoga.

IS: How do you keep going when things get tough?

GL: First, I need to take a deep breath. I like to relax, practicing yoga, sleeping, and meeting my friends and family. The next day I wake up and remember that I am here doing what I like best, and even though I have hard days, I do not want to change course!

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

GL: Wow, a very difficult question. As I can only choose three things: 1) drinking water because without it, nothing would be possible; 2) pocketknife to open seeds, fruits, and whatever else is needed; 3) tent to be safe and to sleep peacefully. If one more item was allowed: lighter to ensure I can have a bonfire! And I hope this island has a lot of food!

IS: Who do you think has had the most considerable influence on your science career?

GL: My doctoral advisor Dr. Sérvio P Ribeiro. He aroused my passion for interaction studies among species and their environments, in search of patterns and their possible explanations. I also have to mention my doctoral co-advisor, Dr. André VL Freitas. He inspired me to dive into butterfly studies, even long before I met him in person. 


Me and my advisors Dr. Sérvio P Ribeiro and Dr. André VL Freitas after my Ph.D. defense (30 November 2018, Unicamp, Brazil)

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

GL: Be aware of the details; field observations are essential for interpreting and discussing the results found! Keep your field notebook well-guarded; it is always useful even after many years.

IS: What is your favorite place science has taken you?

GL: Mexico! During my doctorate, I had the opportunity to spend six months at the Instituto de Ecología – INECOL, working together with Dr. Wesley Dáttilo and his students. For this, I had the support of the University of Campinas and a financing agency in Brazil (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – CAPES, Finance Code 001). It was an unforgettable professional and personal experience. Lots of studying, dedication, and tacos!

In a cannibalistic society, it’s not about survival – it’s all about recycling

A blog post highlighting the article by Chouvenc in Insectes Sociaux.

By Thomas Chouvenc

In this spooky time of the year, there are many examples we could draw from insect societies to give the heebie-jeebies to the non-entomophile. One to usually make it to the news cycle is, of course, the case of zombie-ants. It’s a classic dive into Z-culture while exploring the amazing biology of a host-manipulating fungal parasite. It’s hard to beat as click-bait because it’s just so good. Next in line would be parasitoid wasps, as their gruesome life cycle rips through the host’s organs from the inside while keeping it alive the entire time. Hollywood and manga artists have extensively dug into this concept to bring to life your favorite monster movies and books. Less known, but still a favorite for many of us: the Nicrophorus burying beetle. These Silphidae lay eggs on a decomposing carcass and display extensive parental behavior to their growing larvae as they chew through the putrid flesh of the roadkill. Gruesome, yet full of love: it’s the pinnacle of cute.

So, yes, Halloween could definitely use insects within the spectrum of sociality as core material for horrific displays in our front yards.  With all the classic anatomic fails of course, but this is another horror story. By the way, I was very disappointed when nothing came up from a “parasitoid wasp Halloween display” google search (someone should do something about this, please).

In my opinion, one example that has been either ignored or largely misrepresented in pop culture is insect cannibalistic societies. I will pass on the terrible cliché of old B movies on so-called “cannibal tribes”, but instead bring your attention to termites. Yes, the mostly-ignored or largely-misrepresented group of social insects (wink).  Termites have evolved along with the rise of angiosperms for ~150 Myr, eating a unique niche of untapped dry matter stored in tree trunks that is inaccessible by most plant-feeding animals. Eventually, termites evolved away from their Cryptocercus-like wood roach ancestor and reached the highest level of social organization. They were able to grow large colonies over time while optimizing their ecological success. However, while they enjoyed their pseudo-monopoly on woody material exploitation, their biology was constrained by a significant dietary restriction: wood is carbon-rich but notably nitrogen-poor.


Young and happy termite colony with a healthy king and queen, egg mass, brood, and workforce.

As a consequence, in basal termite taxa, colony metabolism is relatively slow. These taxa rely on their gut microbiota to do the hard work of cellulose degradation. Additionally, they take “forever” to grow, owing to limited nitrogen availability. This whole dilemma has been recognized as fundamentally important for the initial rise of eusociality and subsequent radiation of termites (Nalepa 1984). Termites could obtain all the carbon they needed to fuel the colony by feeding on ubiquitous wood. However, they had to optimize their entire metabolic engine toward nitrogen acquisition, and even more critical: nitrogen conservation. Termites have, therefore, perfected a recycling strategy toward nitrogen conservation over evolutionary time: cannibalism.


Not-so-happy termite colony (Coptotermes gestroi) about to run out of food.

To my knowledge, all termite species perform a colony-wide process of cannibalizing their sick, moribund, and dead individuals. This behavior has received extensive scrutiny by termite researchers around the world because it is so common and evident in termites that one can’t ignore it. Fun fact: when you study the survival of a group of termites in an experiment, you have to count the live termites, not the dead ones, simply because there is rarely a dead body lying around. It is usually cannibalized by nestmates very rapidly.  The concept of “refuse pile”, which can be rather crucial in ants, is mostly useless in termites, as everything goes through someone’s gut, eventually. Cannibalism in termite is, therefore, an essential part of the cycle of nutrients within the colony. It allows precious resources like nitrogen to be reused by the society. This behavior also has attained secondary functions such as sanitation and reducing the risk of diseases within the social group (Chouvenc et al. 2009).

The relationship between cannibalistic behavior and starvation in termites has also previously received some attention. It was suggested that cannibalism can have an essential role in helping a group of termites survive a period of starvation. As soldiers are nutritionally-dependent on workers, they would be eaten in priority. This cannibalism would alleviate their trophic burden to the group and recycle very precious resources toward the rest of the group. Also, the termite group would suppress the production of new soldiers so that only workers would remain, in the hope of restarting the metabolic engine of the group and reducing the trophic burden even further (Su and Lafage 1986). This strategy would, therefore, reduce the metabolic footprint of a colony with finite resources, and aid in more prolonged colony survival.  Soylent green, allons-y!

In a study published online this month in Insectes Sociaux (Chouvenc 2019), I revisited this concept, but this time, I took the “whole colony” approach. Previously, researchers had subjected a small group of foraging termites collected from the field to starvation. To improve upon this method, I subjected 10 whole young termite colonies (Coptotermes gestroi) to starvation. I wanted to take the investigation to a more biologically-relevant scale, with ~3,000 termites per colony, a healthy brood, overlapping generations of workers and soldiers, and of course, the king and queen. The effects of starvation on larger termite groups, the brood, or the primary reproductive were never before investigated.

Recently, I have started revisiting some old questions but scaling things up to the colony level. It can be challenging not to be disappointed in seeing a somewhat different outcome than the one initially expected. To make a long story short: termites are terrible in their survival strategy during starvation events because, in fact, they don’t really have a strategy. Unlike honeybees that store months-worth of honey, termites have a carpe-diem approach to food safety, as they have no internal reserves. Instead, subterranean termites such as C. gestroi will relentlessly (and most of the time successfully) forage for new food sites to prevent food shortage in the first place. But if starvation of the termite colony actually occurs, then the colony just doesn’t have much of a survival strategy. Give it about 30 days.

As the metabolism of the colony is progressively running out of fuel, nutritionally stressed individuals start accumulating in the colony. Unfortunately for termite larvae and workers, who are hemimetabolous insects stuck in a permanent juvenile molting cycle, the time to molt eventually comes, and the younger the instar, the faster the molting cycle. Have you ever tried molting while completely starving?  I would not advise. In my study, this resulted in a failed attempt to molt, death, and subsequent cannibalism from nestmates.

Therefore, the brood and young workers were the first ones to be eaten, not as survival rations for the group, but because they were the first ones to die during their molting process. Then soldiers started running out of juice faster than some of the older workers (who have yet to attempt molting), as they are fed secondhand from workers. Indeed, the remaining workers had to maintain their own metabolism and had nothing left to share with the soldiers, resulting in moribund soldiers, and inherent subsequent cannibalism. In addition, it’s not that the starving colony suppresses the production of new soldiers. It is just that presoldiers that are starving also failed to molt into functional soldiers, and therefore died and were eaten, resulting in an apparent absence of soldier replacement.

Finally, toward the end, a handful of workers remained, with the king and queen being the last ones to die, inevitably.  So, if termites actually have a survival strategy during starvation, it is this: “keep the king and queen alive as long as you can”. Any remaining available energetic resources were eventually funneled to them.

In the end, I found that, contrary to a previous perception, termites don’t reduce the trophic burden of the colony by cannibalizing the dependent castes. Mortality was not cannibalism-driven; instead, cannibalism was mortality-driven.  Termites just do what they always do: if a dead or moribund individual shows up in the group, it is cannibalized to recycle the nitrogen. This is an inherent behavior that was reinforced over millions of years of a nitrogen-deficient diet. The fact is, cannibalizing an energy-depleted individual may not provide much energy to the group, as you can’t recycle ATP that does not exist.

The excessive mortality resulting from starvation triggered a massive cannibalism wave, which ended up with an accumulation of old workers with staggering levels of uric acid building up in their fat body. Such an observation is typical of termite colony collapse, as there is a sudden excess of available nitrogen through cannibalism, something termite metabolism never evolved toward.  By 20 days after the starvation was in effect, the colony started shutting down, and cannibalism was no longer observed. Dead bodies began accumulating in the colony and were not taken care of by the surviving workers, which were focusing only on the king and queen. In the end, the king and queen eventually starved to death too. So much for an efficient survival strategy in termites.

In this study, despite the amount of work it took to investigate a question at the colony-level and provide detailed observation, it was, in my opinion, absolutely worth it. Experimenting on various aspects of termite biology in small groups in a Petri dish can provide valuable initial information. However, scaling it up to a colony-wide observation can drastically change our understanding of a social group.  For example, if you have a termite infestation in your house, you can now actually say “my house is being attacked by a horde of cannibals.” It would be almost accurate. Happy Halloween.


Although this paper discussed how cannibalism can go wrong in a starving termite colony, this picture reveals a case of cannibalism gone (very) weird – if one can make any sense out of it. It depicts a worker in the process of cannibalizing a nestmate, but his head is stuck inside the terminal abdominal segments of the dead one. Photo courtesy from David Mora Del Pozo.

Thomas Chouvenc is an Assist. Professor in urban entomology at the University of Florida, UF/IFAS.

Twitter: @ChouvencL


Chouvenc T (2019) Limited survival strategy in starving subterranean termite colonies. Insectes Sociaux.

Chouvenc T, Su NY, Robert A (2009) Inhibition of Metarhizium anisopliae in the alimentary tract of the eastern subterranean termite Reticulitermes flavipes. J Invertebr Pathol 101: 130-136

Nalepa CA (1994) Nourishment and the origin of termite eusociality. Nourishment and Evolution in Insect Societies (ed. by: J.H. Hunt & C.A. Nalepa), pp. 57-104. Westview Press, Boulder, Colorado

Su NY, La Fage JP (1986) Effects of starvation on survival and maintenance of soldier proportion in laboratory groups of the Formosan subterranean termite, Coptotermes formosanus (Isoptera: Rhinotermitidae). Ann Entomol Soc Am 79: 312-316

Interview with a social insect scientist: Claire Morandin


IS: Who are you and what do you do?

CM: My name is Claire Morandin, and I am a postdoctoral researcher on an EMBO fellowship at the University of Lausanne, but currently on maternity leave – my second daughter was born in July. When I am not taking care of her, I am mainly spending my time conducting research that focuses on social insect genomics. In general, I am interested in understanding how complex phenotypic traits arise and evolve across species. And more specifically, I study the evolution and maintenance of the female castes in ants. Recently I started working on honey bees, too, trying to answer similar questions in the evolution of female castes. I got hooked on bioinformatics during the work for my Ph.D. thesis, and I use genomics tools such as RNA sequencing, comparative genomics, evolutionary analysis, gene co-expression networks, and methylation to understand these complex mechanisms.

IS: How did you develop an interest in your research?

CM: After finishing my bachelor’s degree, I took a year off and volunteered with the CNRS to work in Zimbabwe, basically being a field assistant helping scientists with their experiments. Having had this experience, I discovered my passion for research and decided that I also wanted to conduct my own research experiments and become a scientist myself. This decision led me to continue with a master’s at Uppsala University in Sweden and a Ph.D. degree at the University of Helsinki in Finland. It took me a few more years of studying and meeting the right mentors to discover the exciting world of social insect research.

IS: What is your favorite social insect and why?

CM: Before last year I would have definitely answered ants, probably Formica ants. However, honey bees are cool too and way easier to study. They can develop from eggs to adults in the lab, and I am fully taking advantage of this feature at the moment. So, I would go for honey bees being my favorite social insects now. Also, as a side product of keeping bees, you get plenty of honey!

IS: What is the best moment/discovery in your research so far? What made it so memorable?

CM: The best moment was my Ph.D. defence; it was challenging but also a lot of fun to wrap up four years of hard work and present what I had accomplished during these years. This process makes you realize all the smaller puzzle pieces coming together and revealing new bigger conclusions. I also had a great discussion with my opponent, Dr. Seirian Sumner. You do not know how enjoyable a defence is before you get the chance to have such a long and stimulating discussion with a great opponent. Also, having all my family and friends and people who encouraged me through the years together in one room to celebrate with me made this day so very special (and well, the party afterwards was pretty great too!).

IS: Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

CM: I sadly do not do much outreach, besides communicating my own and research I find generally interesting and exciting over social media. However, I occasionally teach bioinformatics and try to spread my enthusiasm for this field to the students. I have also recently gained an interest in gender studies. With a colleague, Dr. Luke Hollman, we recently published a paper showing that researchers tend to collaborate with same-gendered colleagues leading to further discrepancies between genders in life science. I think this is an important and very relevant topic in science, although not related to my actual research. With such insight, we can raise awareness of the gender gap in research, which eventually may contribute to making this gap smaller and hopefully disappear in the decades to come.

IS: What do you think are some of the important current questions in social insect research and what’s essential for future research?

CM: There is so much we still do not understand about social insects, and that makes my work very exciting. Despite plenty of social insect genomes out there, we are still trying to put pieces together to determine whether common mechanisms are behind caste differences across all social insects. More comparative genomics work will be essential to shed light on such important evolutionary questions.

IS: What research questions generate the biggest debate in social insect research at the moment?

CM: I would think that some of the biggest questions are the ones related to the origin and evolution of eusociality across the phylogeny tree. However, from my current work point of view, I would say that the role of methylation in caste differentiation also generates lots of debates that I like to fuel up with more data and findings from my work.

IS: Outside of science, what are your favorite activities, hobbies or sports?

CM: I love running and baking, and recently took a significant interest in sewing – it helps me calm down when everything else around me spins so fast.

IS: How do you keep going when things get tough?

CM: When things get tough, I like to go outside for long walks or runs with very loud music to forget about everything. And then I come home to two happy daughters who only want to play and laugh, and I quickly forget about the tough things because making them happy is what matters most. Having a life outside of science – a place to go when things get tough – helps me go through the stress and the hardship of a scientist’s life.

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

CM: A machete, a water bottle and a book (would finally have time to read a grown-up book, and not just children’s books).

IS: Who do you think has had the most considerable influence on your science career?

CM: My collaborator and mentor Sasha (Dr. Mikheyev) has had the most considerable influence on my science career. He made me go from qPCR to RNAseq, and I never looked back. Because of him, I got hooked on bioinformatics.

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

CM: Be patient – very patient – and do not forget that science is fantastic but that it is not everything in life, so when it gets tough, you always need something else that can make you happy. That was more general advice for being a scientist, I guess. For social insects, there is not much to say. They are fantastic, go for it!

IS: What is your favorite place science has taken you?

CM: Japan, I have had the chance to visit Sasha’s lab at the Okinawa Institute of Science and Technology (OIST) eight times already, and I love going back there every year. Okinawa is so beautiful!

Interview with a social insect scientist: Edith Invernizzi


IS: Who are you and what do you do?

My name is Edith Invernizzi, and I am a Ph.D. student in the School of Biology at the University of St Andrews (Scotland), studying collective behaviour and self-organisation. I am a theoretical modeller, mainly, but I believe that all models should be built in a constant feedback loop with real data. I try to integrate laboratory and field experiments in my work as much as possible.

IS: How did you develop an interest in your research?

Since I started working on models for an undergraduate project, I have been interested in the evolutionary dynamics of behaviour. It’s this that ultimately led me to collective behaviour and ants in my Ph.D.

IS: What is your favorite social insect and why?

Ants, because of the immense ecological range this genus has been able to colonise. They are a perfect case study for different species that can independently evolve similar collective behaviour mechanisms when the environmental context is different.

IS: What is the best moment/discovery in your research so far? What made it so memorable?

I’m very early on in my research, so I have a limited range of experiences to draw from. But I do know what the best moment has been. When I started looking at social insect collective nest building, I came to the hypothesis that self-organised activity could be seen as continuous assessment of the environment, a homeostatic mechanism in response to constant fluctuations. I then discovered the body of literature that proposes precisely that frame of interpretation. I felt that I was learning how to understand behavioural data and, most of all, that my way of understanding was meaningful to other scientists.

IS: Do you teach or do outreach/science communication? How do you incorporate your research into these areas?

I personally do not do much outreach, but I try focusing on how social insects make collective decisions whenever asked about my research by a non-scientist. I think decision making is a fascinating behaviour that every human can relate to, but that is often considered only from an individual perspective. I try to use this example to give a different point of view.

IS: What do you think are some of the important current questions in social insect research and what’s essential for future research?

The importance of individual-level variation when we think of colonies and eusocial species. There are suggestions that behavioural variation within a colony causes different individuals to exploit different resources. Similarly, in behaviours such as collective building, workers with varying levels of susceptibility to different cues might initiate changes in the structure that are important for short-term adaptation. I suspect that understanding the role of within-group variation is not only crucial for helping disentangle environmental adaptation but also has potential applications in fields such as robotics and engineering – for instance, in Ant Colony Optimisation (ACO) algorithms.

From a ‘genetics versus plasticity’ perspective, it is interesting to study how much of this variation derives from allelic genetic differences, in poly-mated queen or multiple-queen colonies, and how much from epigenetic factors. For intra-colony competition, the question becomes evolutionary: is there an ‘optimal’ level of inter-individual variation that maximises colony fitness? How might mating systems, genetic and epigenetic mechanisms have evolved under selection to reach and maintain such level?

To start scratching the surface of these questions, we need more tools that facilitate individual identification and individual behaviour records. Developing more advanced movement and ethological tracking methods (the latter automatically tracking behavioural events in a video recording), for example, is essential if we want to obtain a large amount of data with reliable sample sizes and make this fine level of individual detail in group behaviour studies the norm. More behavioural studies done in the field are also necessary to match the observed variation to the multiple factors that it might respond to, in a complex environmental context.

IS: What research questions generate the biggest debate in social insect research at the moment?

Probably the questions about cognition. How advanced are insect cognitive capacities? Are we underestimating them or overestimating them? What is the relationship between individual cognition and collective cognition at eusocial colony-level?

IS: What is the last book you read? Would you recommend it? Why or why not?

A Time for Everything, by K.O. Knausgaard. If you are into rational psychological analysis and minute behavioural details but still like to find an overarching narrative when you look at life experiences, then Knausgaard is the author for you.

IS: Outside of science, what are your favorite activities, hobbies or sports?

I practice martial arts, read contemporary literature, and enjoy planning exciting activities. I prefer a good stand-up comedy show, live or on TV, over a holiday trip anytime.

IS: How do you keep going when things get tough?

When nothing seems to work, I remind myself that science and my Ph.D. are only one aspect of my life, and I try to put more energy into others. If I meet a wall in my research, then I create something else or break down a barrier elsewhere. Working on other projects just remind me that I can produce an output I am enthusiastic about and renew my creativity.

In tough times, it is focusing on what I find exciting about the work, the goal, or the challenge that keeps me going. But I must schedule some fun activities throughout the week that break the tension and give me fresh energy; a good climb or stand-up comedy always do the job.

IS: If you were to go live on an uninhabited island and could only bring three things, what would you bring? Why?

A Swiss knife, a water flask, and a diary with a pen attached to it. The first two for surviving and the third one to keep my sanity. I enjoy spending time alone, and if I have the time, I might as well keep track of my thoughts.

IS: Who do you think has had the most considerable influence on your science career?

Definitely my undergraduate supervisor and master’s co-supervisor, R.T. Gilman. He has given me a passion for dissecting dynamics, and he is the one who introduced me to modelling.

IS: What advice would you give to someone hoping to be a social insect researcher in the future?

Be patient. Be passionate. And always work with a behavioural scientist if you are a theoretician. They bring you down from your matrices into the real world.

IS: What is your favorite place science has taken you?

I did my Master of Philosophy fieldwork in the area around Lugu Lake, in South Western China. It is close to the most beautiful place I have ever been to, and I hope that its landscape and culture have remained unspoiled.