Interview with social insect scientists: Kaleigh Fisher and Mari​ West

Kaleigh and Mari are authors of the recently published review article, ‘Are societies resilient? Challenges faced by social insects in a changing world’.

 

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Kaleigh Fisher doing fieldwork in Chiapas, Mexico

 

 

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Mari West

 

IS: Who are you and what do you do?

KF: I am a graduate student in the Woodard lab at the University of California, Riverside. I am using methods from insect behaviour, evolution, and sensory ecology to understand how taste operates in bumble bees.

MW: I am a 3rd-year graduate student in the Purcell Lab in the Entomology Department at the University of California, Riverside. I study non-reproductive division of labor in ants, with the goal of understanding how continuous size variation and differences in chemical cues among workers affect how tasks are partitioned in social insect colonies.

 

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Kaleigh Fisher doing fieldwork in Chiapas, Mexico

 

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

KF: I was interested in agroecology and started working in a lab that studies insects in shade coffee farms in Mexico. I became interested in the ants and the stingless bees in this system. The research was super exciting; it was my first introduction to social insects, and I have been working with them ever since.

MW: I first became interested in social insects as an undergraduate at Cornell University, where I worked in Dr. Linda Rayor’s lab, studying behavioral dynamics between reproductive females in social spider colonies. These spiders are usually very tolerant of each other but can become quite aggressive (to the point of being cannibalistic) when vying for reproductive opportunity. Later, while working with an invasive ant species that is highly polygyne and super-colonial, I realized that there was a vast spectrum of cooperative behaviors within social groups. Since this realization, I have been interested in how social insects can coordinate these collective behaviors without any central control.

IS: What is your favorite social insect and why?

KF: My favorite social insects are the stingless bees. They are so interesting both biologically (eusocial, ~500 species) and culturally (long history of stingless beekeeping in Mexico). I worked with them briefly before starting to work with bumblebees and hope to have the opportunity to work with them in the future.

MW: Why do I have to have a favorite? They’re all awesome in their own ways.

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

KF: I think the best moment in my research so far happened a few weeks into my first field season on a shade coffee farm in Mexico for my master’s research. I stopped and thought, “wow this is awesome, I can’t believe this is my job!” It’s easy to get overwhelmed by all of the components involved with being in academia; I think it’s important to remind myself from time to time how exciting it is to ask a question about why or how insects are doing something, especially in the field, and then figure out a way to answer it. I realized how awesome that is in this moment. 

MW: Before I started my graduate work, I volunteered for a research project with the US Fish and Wildlife Service’s Crazy Ant Strike Team, whose goal is to eradicate the invasive yellow crazy ant from a Pacific island. The island serves as important nesting habitat for ground-nesting seabirds, which the ants significantly disrupt. My crew’s goal was to test a few different control/eradication strategies so that the next team could implement the most effective one. The project has been very successful, leading to almost complete eradication of the ants and a significant recovery in seabird reproduction on this island! It feels incredible to have been involved in a conservation project that has made a positive and lasting impact!

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

KF: Yes, to both; I enjoy teaching (I have only had the opportunity to be a teaching assistant so far) and doing outreach. Depending on what I am teaching, I try to use relevant examples from social insects, especially bumblebees. I also really enjoy outreach. Depending on the group, I will introduce my specific research and why I do it or discuss the importance of pollinators and insects in general.

MW: Both as an undergraduate and graduate student researcher, I have been lucky to be part of entomology departments with strong focuses on outreach and I have been involved in many large-scale insect fairs throughout my academic career. Additionally, I have made it into many K-12 classrooms through these programs. I always try to incorporate my research into outreach events by introducing the basics of the colonial lifestyle and cooperative behavior of social insects. For more advanced audiences, I will also explain my experimental designs and findings.

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

KF: A question that I am interested in is how environmental context shapes social behaviour. A lot of super informative research about insect sociality has come out of laboratory studies. I think building on those findings to capture how variable they are across different populations and species is essential.

MW: If you read our paper, Fisher & West et al. 2018, then you already know. 🙂

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

MW: My general feeling is that a lot of debate surrounds honeybees – whether or not they are good/necessary for ecosystem functioning and whether their long history of being farmed makes them a useful model system for studying social insects.

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

KF: The last book I read was the Ministry of Utmost Happiness by Arundhati Roy. It is fiction, but it has powerful social and political commentaries about current events in India and globally. I enjoyed it, so I would recommend it to those who enjoy novels with strong social commentaries.

MW: The Jungle, by Upton Sinclair. It’s certainly not very upbeat, but I do think it is relevant to some social and economic issues that our world faces today.  It was a thought-provoking read.

 

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Kaleigh Fisher and family in Chiapas, Mexico

 

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

KF: Outside of science, I like to spend time with my family (hiking, gardening, cooking together).

MW: Hiking and baking. I like exploring the outdoors, mainly to observe wild animals in their natural environment, and working with my hands to make something tasty to share with others.

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

KF: Whenever I feel overwhelmed, I try to step away from everything for a moment, even if that means just going for a walk, so that I can get a better perspective on everything.

MW: I usually remind myself how lucky I am that my job brings me outdoors on a regular basis and keeps me intellectually engaged. I try to give myself some time to relax outside, to reflect and refresh my outlook on life.

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

KF: I would bring my partner because he is an expert field biologist and awesome human, and a notebook and a pen (to help organize my thoughts/ideas).

MW: Having previously lived on an uninhabited island to which I was able to bring pretty much everything I ever wanted, this is a tough question to answer. However, if I had to choose, I would bring a snorkel for exploring the coral reefs (and maybe making fishing a little easier), some sunblock, and someone with whom to share the experience. In my experience, you don’t need much more.

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

KF: I think my master’s thesis advisors have had the most significant influence on my scientific career. They have inspired me to be inherently curious about insects while simultaneously instilling in me the importance of doing science consciously and with a moral compass.

MW: Without Dr. Linda Rayor’s influence, I probably wouldn’t be studying social insects today. She was an incredible mentor, and her enthusiasm for and ability to communicate about her science caught my attention immediately. In addition to getting me interested in social insect behavior, she has inspired me to share my science with others around me.

 

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A bumblebee foraging experiment conducted by Kaleigh Fisher

 

 

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Ant abdomen painted by Mari West during a mark-recapture experiment

 

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

KF: Give yourself time to explore what kind of research you are passionate about. Being excited about the questions you are asking and the research you are doing is what it is all about.

MW: I would encourage them to spend some time working on different projects so that they can identify the questions that they are most excited about and learn about what kinds of experimental design work well for those questions. I would also encourage them to spend many hours watching their study organisms, for fun and inspiration.

Interview with a social insect scientist: Thiago Silva

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IS: Who are you and what do you do?

TS: My name is Thiago Silva, and I am a researcher at the Universidade Federal do Paraná, Brazil. During my Msc. I studied the diversity of two genera of myrmicine ants (Acanthognathus and Strumigenys) in the Brazilian Atlantic Forest. My Ph.D. thesis stretched a bit to the morphological side, and I explored some aspects of the anatomy of several species of Strumigenys using web-based tools to annotate classes in online anatomic ontologies.

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

TS: As an undergraduate student, I became obsessed with insects, mostly wasps and grasshoppers. At the time, my supervisor (who is an ornithologist) was studying the edaphic fauna of one of the few national parks we have at the southern region of the Atlantic Forest. Knowing of my interest in insects, he invited me to study the ant fauna he collected at the site. With some reluctance (I really loved wasps) I accepted the challenge. When I laid eyes on my first mounted ant under the stereoscope, it was love at first sight.

IS: What is your favorite social insect and why?

TS: I love minute and shy social insects like Strumigenys inusitata. It is a bizarre-looking ant with a flattened head and a distinct hump over its mandibles (see image below). We do not know what the function of this structure is, but since most species belonging to this genus are slow-paced hunters living at the leaf-litter, we guess it might be related to a prey attraction strategy.

 

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Scanning electron microscopy of the head of Strumigenys inusitata. Left is anterior. Note the distinct hump at the anterior margin of the head.

 

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

TS: I do not remember any specific great discoveries during my research. I think that seemingly-small everyday discoveries, such as learning a little bit about the morphology of a cool group, are the most memorable moments to me.

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

TS: I often talk about science and biology to undergraduate and graduate colleagues, whether in more casual settings at the university or during courses. I always try to highlight the importance of morphology to other disciplines in biology and how the study of phenotypes provides all sort of hints that enables us to understand a lot about biological systems.

Recently, I started talking more about representation of knowledge and the importance of clear communication in science. It is a somewhat new topic for me, so I am gradually incorporating these ideas in my talks.

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

TS: Since social insects are extremely complex, I think there are numerous important questions that have to be (and are being) addressed. I believe that the most essential one is how sociality came to be in different groups of insects. Although its partial understanding does not impede the exploration of other important aspects related to sociality, I think the complete understanding of the origin of sociality is a primary topic of research for social insect scientists worldwide.

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

TS: Terminology always strikes me as the hot topic in my field of research and I think I always focus on it when reading about social insects.

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

TS: The last non-fiction book I read was Bruno Latour’s We Have Never Been Modern. It is a fascinating book and I recommend it, especially in the current social and political global context. The book deals with the modern duality of culture and nature and how this polarity affects decision-making at different levels within modern society.

The last fiction books I read were Haruki Murakami’s Kafka on the Shore and the graphic novel Black Dog: The Dreams of Paul Nash by Dave McKean. I would recommend any of the Murakami’s books, but Kafka is a favorite of mine since the characters depart from the typical constructs the author used in his previous works. Also, I love Murakami’s magical realism and references to popular culture. Black Dog is a must-read for those who want to take a glimpse at the nightmarish effects of war and understand how the first world war affected (and still affects) people at local and global scales. The mixed painting styles used by McKean and how their interplay with the narrative makes the reading extremely joyous.

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

TS: I spend most of my time outside of academic grounds reading or writing. I also like to get on the road and travel to other places. Sometimes one’s own mind is the best hobby one can have.

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

TS: I write my concerns away. I discovered that it is a useful sort of therapy for me. It organizes my thoughts, clearing the path that leads to the most unstressful resolution of a problem.

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

TS: Paper, pencil and the opportunity to leave the island! As Donne said, ‘every man is a piece of the continent’. Although I am very fond of reclusion from time to time, it is nice to be with kin.

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

TS: My parents and brother. My mother is a former researcher in obstetrics and always studied postpartum wellness and scientific methodology. My brother is an archaeologist/anthropologist and has always been concerned with the relation of public patrimony (especially archaeological sites) and traditional communities. My father is not a researcher, but always shared with me the delights of investigation and discovery. I think that a person’s major influences are those that are closest to them, providing support and teachings wherever they go.

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

TS: Love what you do, try new things, and don’t be afraid to make mistakes.

Interview with a social insect scientist: Joel Woon

 

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Photo credit: Matt Jarvis

 

IS: Who are you and what do you do?

JW: My name is Joel Shutt Woon, and I have just started a Ph.D. at the University of Liverpool, where I plan to research climatic tolerances of termite communities in Western Africa. However, as with all research projects, the fine details could change! Before this, I studied at Imperial College London for both my BSc in Zoology and MRes in Tropical Forest Ecology.

 

termites are working on tree bark. unite team work  for harmonious working.

Photo credit: Adobe

 

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

JW: I was lucky enough to grow up between two large parks in Sheffield, UK, which gave the perfect opportunity to immerse myself in nature. I’d spend hours exploring the forests, searching for insects and amphibians, or following bird calls to try to catch a glimpse of the culprit. This experience growing up developed into a love for the natural world and made my course choice at university very simple. At university, through my BSc and MRes, I found that tropical rainforests were the habitat that enticed me the most. The ability to study your passion in some of the most incredible environments in the world is extraordinary and something that I hope to do for a very long time. I have also discovered, mainly through the course of my MRes work, that I am captivated by entomology, and specifically termites. Their importance as ecosystem engineers, nutrient cyclers, pests, and a food source makes them fascinating to study, and to top it off I think they look charming, although I accept I might be alone in that.

 

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Photo credit: Adobe

 

IS: What is your favorite social insect and why?

JW: There are so many cool termites to choose! Globitermes globosus has a remarkable defensive strategy that involves biting an attacker, locking its jaw, and then secreting a sticky liquid out of its head. This causes the termite to ‘glue’ itself to the attacker (usually a pesky ant), immobilising it and giving time for other soldiers to arrive and workers to repair the nest. Hospitalitermes march through rainforests in groups, similar to army ants, and you can see where they’ve been after they’ve disappeared because all the wood in their path will have been cleared of microepiphytes, leaving a clean trail. Moreover, you have to admire the savannah species of Macrotermesfor the complex and massive nest structures they manage to build.

 

Cordyseps2 Credit Matt Jarvis

Photo credit: Matt Jarvis

 

Away from termites, Camponotus gigas (giant forest ants) are exciting to see around the forest; true gentle giants of the insect world! The size of them blew my mind when I first saw them. I also saw one infected with a cordyceps fungus – seeing a fungus growing out of a giant ant was like witnessing science fiction come to life!

 

african landscape with termitary

Photo credit: Adobe

 

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

JW: I’m still in the fledgling stages of my research career, so I haven’t had many “discoveries”, but finishing my first thermal tolerance experiment was an exciting moment. My supervisor had carried out the same tests on ants, so we started with low expectations of termite thermal tolerance and those expectations were exceeded! Also, walking out into the field to work on my own project for the first time was a huge moment for me. It was the realisation of a goal and a lot of hard work!

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

JW: I don’t currently do outreach, as I am in the opening week of my Ph.D. as I write this, however, it is something that I want to develop. One of the biggest challenges we face as scientists is communicating our research to a broader audience. We can affect much more significant change if we communicate well than if we keep our work isolated within the scientific community. I think it is an area of science that needs a lot of improvement, and I want to contribute to that improvement.

 

Namibia termite mounds

Photo credit: Adobe

 

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

JW: We need to further our understanding of how climate change will impact social insects and increase the power and accuracy of these predictions. Social insects are incredibly important to a myriad of ecosystems, in a plethora of different ways, so understanding how a changing climate will impact their roles within those ecosystems is essential. Will climate change cause species distributions to change? If social insect species are displaced from ecosystems, is there redundancy to cover the lost ecosystem services? How will social insects impact the ecosystems to which they get displaced? These questions remain important.

 

Macro of termites on the forest floor, Borneo, Malaysia

Photo credit: Adobe

 

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

JW: Quantifying and understanding the ecosystem services social insects provide is still a huge topic. Because many species of social insects are massively abundant, they can make huge impacts on ecosystems worldwide. Quantifying those impacts, and what happens to those ecosystems when social insects are excluded, is extremely important. It not only allows us to understand the relative importance of species, but also apply ecological and monetary values to them (regarding ecosystem services and damage caused), which is extremely useful in our modern world.

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

JW: I just finished the Conclave of Shadows saga by Raymond E. Feist, which is the fourth saga set in the fictional world of Midkemia. I would recommend reading Feist’s first book, The Magician, to all fans of fantasy, as it is an excellent and archetypal mythical story that many modern classics borrow from. I’m also reading The Hidden Life of Trees by Peter Wohlleben, which is a fascinating popular science book for people like me who haven’t had much experience in dendrology.

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

JW: My main passion, aside from research, is travelling. I love immersing myself in new cultures, having new experiences, and seeing new environments. Through travelling, I’ve been lucky enough to visit some of the most amazing areas for marine life in the world, which has fostered a passion for scuba diving. Scuba diving is a very surreal and exhilarating experience and one I would recommend everyone try at least once. There is nothing else like it. When I’m back in the UK, my main hobby is playing a card game called Magic: The Gathering. It’s incredibly exciting and complex (it holds the world record for the game with most rules!) and has so much variation while challenging the player to think and strategise well.

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

JW: I think it’s essential to have a dependable support network for you to fall back on. Whether that’s scientific support or emotional support, having people who you can talk to openly and honestly and rely on for help allows you to work through all sorts of issues. I also think you need to have a hobby that you can step away to. Whether it’s for an hour, a day, or a week, you need something that is unrelated to research that you can occupy yourself with, and that will allow you to diffuse any negativity towards your work.

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

JW: I would take an e-reader filled with books (including multiple on how to survive in the wild), a solar charger, and a satellite phone to ring for help when something inevitably goes wrong.

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

JW: As with many biologists of my generation, the person who started me down the path I’m on is Sir David Attenborough. His documentaries opened my eyes to the incredible diversity and majesty of nature and made me want to pursue a career studying it. In a more practical sense, Mike Boyle, my friend and MRes supervisor, has also contributed massively to my development as a scientist, providing invaluable support, advice, and training as well as a healthy dose of realism.

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

JW: Be inquisitive and ask questions, and follow what you find interesting. Scour the internet/ library, go bug hunting in a garden, or contact experts! Most scientists love to share their research; whether you’re a grade school student or undergraduate, an amateur or a professional – if you reach out to people in your community, it is incredibly likely that you will get a response and tap into a source of knowledge that you wouldn’t otherwise be able to access. So send an email or two, ask a few questions, and see where it takes you!

Honey Bee Immunity: More Specific than We Thought

A blog post highlighting the article by N. Wilson-Rich, R. E. Bonoan, E. Taylor, L. Lwanga, and P. T. Starks in Insectes Sociaux

By Rachael E. Bonoan and Philip T. Starks

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Nature can be hostile, especially for insects — tiny creatures that deal with big problems. They get eaten, they dry out, they get smashed. Believe it or not, insects also deal with even tinier pests and pathogens, just as we do.

Also, like us, insects have developed an immune system to combat such microscopic threats. Over the years, scientists have uncovered how insect immunity relates to behavior, mating success, ability to find food, nutrition, energy cost, etc. However, the method used to study insect immunity — sterile fishing line inserted through the membrane between the sclerites — does not reflect the evolutionary history of insects and their pathogens. In our recent paper, we describe a modified method to investigate insect immune strength and its specificity. We show that the insect immune system may be able to recognize different classes of pathogens and respond accordingly.

An insect’s first line of defense is a physical barrier: the exoskeleton. When the exoskeleton is injured, however, diverse pathogens can invade. Once a pathogen makes it past this barrier, the insect immune system uses proteins called pattern recognition receptors (PRRs) to recognize cellular patterns on the invader (pathogen-associated molecular patterns, PAMPs). One of the many responses the immune system can then carry out is the encapsulation response (ER) where the invader is surrounded by immune cells which secrete a protein, melanin, that detoxifies the invader.

The strength of insect ER is typically measured by inserting a piece of sterile fishing line into the insect, waiting for the immune system to respond, and then removing the fishing line “invader.” Researchers then use light microscopy to measure how much melanin (i.e., color) developed on the fishing line. The darker the explanted fishing line, the more melanin deposited, and thus, the stronger the immune response. While the fishing line does mimic a physical injury, it does not mimic the diverse pathogens with which insects have evolved.

To make this method more evolutionarily relevant, we added a layer to the typical method: PAMPs. Different pathogens have different patterns (PAMPs) on their surface that the host PRRs recognize. We tested honey bee ER in response to fishing line coated in PAMPs found on two types of bacteria and fungi. We call our modified fishing line implants PAMPlants.

Once we coated the PAMPlants, we carefully inserted them between two segments of the honey bee exoskeleton. This is truly a labor of love. Handling the coated fishing line (2mm long, 0.4mm diameter—tiny!) with forceps takes steady hands and attention to detail. Thankfully, two NSF Research Experience for Undergraduate fellows were around to help carry out this mini-surgery on the 176 bees it took for this study! To let ER happen, we left both uncoated implants and coated PAMPlants in the honey bee for 4 hours. We then carefully removed all implants, and prepared a microscope slide with the explant. Microscopy was used to take images of the explant. We analyzed the images for color (i.e., melanin) as the typical proxy for ER strength.

Compared with a control implant (fishing line coated in phosphate-buffered saline), honey bees had a stronger ER to PAMPlants. In honey bees implanted with both a control implant and a PAMPlant, fungal PAMPlants and one of the bacterial PAMPlants (lipopolysaccharide) lead to a stronger ER than the control implant.

With this modified method, we show variation in honey bee immunity in response to different classes of pathogens. Since we saw an increase in ER for fungal and one of the bacterial PAMPlants, it is likely that physiological immunity is important for fighting these types of pathogens in honey bees. We did not see similar results for our third PAMPlant (peptidoglycan) which is found on some bacteria. In agreement with our results, honey bees likely combat this specific type of bacterial infection behaviorally rather than physiologically (Spivak and Reuter 2001).

When it comes to dealing with pests and pathogens, honey bees have it especially hard. In keeping bees, humans have unwittingly facilitated the spread of honey bee disease. In commercially raising bees, we have increased the density of honey bees across the landscape, and in some cases, we nurse weak colonies which are more likely to spread such disease.

The most notorious honey bee pest is the Varroa mite. This mite is the honey bee’s version of a tick. Ticks spread diseases in humans by puncturing our protective barrier (i.e., skin) and feeding on our blood. Varroa mites spread disease by injuring the honey bee’s physical barrier and feeding on the bee’s fat body—an essential organ for energy storage and immunity (Burnham 2018; Kielmanowicz et al. 2015).

Our modified method suggests that uncoated implants (i.e., fishing line) may not give insect immunity enough credit. While we have learned a lot about insect immunity with uncoated fishing line, we have so much more to uncover with PAMPlants!

References

Burnham T (2018) Downtown new hope in the fight against Varroa. Bee Culture, A.I. Root Company, Medina, OH, USA

Kielmanowicz MG et al. (2015) Prospective large-scale field study generates predictive model identifying major contributors to colony losses PLoS Pathog 11:e1004816 doi:10.1371/journal.ppat.1004816

Spivak M, Reuter GS (2001) Resistance to American foulbrood disease by honey bee colonies Apis mellifera bred for hygienic behavior Apidologie 32:555-565

Information choices for navigation

Highlighting the article written by Middleton, Reid et al. in Insectes Sociaux

Written by Insectes Sociaux Editor-in-Chief, Michael Breed

What happens when an animal has choices of navigational information from two or more sources? In foraging social insects, this choice often occurs when workers have the decision to use either learned navigational cues and pheromone trails or markers. Using learned cues may involve trade-offs between speed and accuracy, as movements oriented to landmarks are usually faster than movements based on continuous search for pheromone cues, but the workers may be less prone to errors when following pheromone trail.

An added wrinkle to this trade-off is that the accuracy of landmark based navigation usually improves with experience. An animal that has navigated a route several times may move more confidently, accurately, and rapidly than a naïve individual. This experiential shift can also represent a gradual shift from the use of social cues, such as pheromones, to internalized landmark memory (which can be considered private information), with some reliance on both types of cues during intermediate stages.

In this issue, Middleton, Reid, and their colleagues investigate these shifts in information usage in Australian meat ants, Iridomyrmex purpureus, using a y-maze experimental design (Middleton, Reid et al 2018). They found that, as with other species of ant, experienced workers use landmark information in preference to trail pheromone information, and that if the trail information is removed, experienced workers continue to be able to navigate the route. The memorized route information is considered private (internal to the individual) while the pheromone information is public (available to all colony members).

A unique element of this study is the discovery that the meat ant workers’ performance using private information does not improve with repeated experience; they do well with private information after just navigating the route once. Regarding learning style, this rapid acquisition of information matches well with imprinting, in which an animal rapidly acquires critical information. This pattern differs from the learning curve, showing improvement over trials, that typically results from trial and error learning. This finding supports the critical role of navigational accuracy in successful social insect worker foraging. It would be fascinating in future studies to further explore the speed with which navigational route is acquired by ants and to compare those results with the much more thoroughly studied honey bee.

Because foraging by social insects is usually a cooperative, rather than a competitive, venture for workers from the same colony, the concept of private versus public information applies differently than in other behavioral contexts. In systems involving competition, including attraction of mates or food search by non-cooperating animals, information is held privately because it has particular value to the holder, and that value would be compromised if other animals, by eavesdropping or spying, capture the information.

The use of the public/private categorization in this context, though, is intriguing because it raises the question of at what point would a forager become motivated to share its private information publicly, by re-laying pheromone trail or communicating an alternative more direct route to the goal? This question leads to additional possible future directions, with studies focused on the mechanisms of choice between continuing to forage and shifting to providing social information to colony-mates.

References

Middleton EJT, Reid CR, Mann RP, Latty T (2018) Social and private information influence the decision making of Australian meat ants (Iridomyrmex purpureus). Insect Soc
doi.org/10.1007/s00040-018-0656-1

Interview with a social insect scientist: James Glasier

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IS: Who are you and what do you do?

I am an instructor at the University of Alberta Augustana. I did my BSc and MSc at the University of Alberta, Canada, and recently graduated with my Ph.D. from the University of New South Wales, Australia. Most of my research has focused on ant ecology and other myrmecological topics. A lot of my research has focused on ant diversity and ecology in western Canada. However, I also developed an interest in the global patterns of myrmecophiles (organisms that closely associate with ants) during my Ph.D. studies.

 

iridomyrmex purpureus

Iridomyrmex purpureus

 

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

Well, I have been interested in ants ever since I was a kid playing in the garden; I would sit in the grass and just watch them do what ants do. I also like the thrill of discovery, and at one point in my life I wanted to be a palaeontologist because I loved finding fossils and discovering something that no one had ever seen before. Science, however, is often about discovery and that feeling of wonder can be found in any scientific topic. During my undergraduate degree, I did an independent project in entomology, specifically looking at ants from Alberta. This project captivated me, as it seemed like every time I looked under the microscope I was finding a new ant record for the area. Finding out that we knew so little about ants in western Canada led me to switch from palaeontology to entomology and I have been studying ants ever since!

 

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Lasius neoniger

 

IS: What is your favourite social insect and why?

That is a difficult question, as there are so many fascinating social insects. I would have to say Formicoxenus ants are some of my favourites. They are social parasites of larger ants, such as Formica or Myrmica, and live within their host ant colony. These much smaller ants use their hosts for protection, but also steal food from them by running up and begging to be fed. Moreover, the male ants of these species have evolved to be more worker-like, often are wingless, and may actually help in colony activities! These unique ant traits make Formicoxenus a fascinating genus of social insects.

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

Probably discovering that there are a lot more ants in Alberta than was previously thought. When I started my work, there was believed to be about 50 ant species; now that estimate is closer to 100, not including introduced species. For some ants, their known range was extended by over 1500 km. For me, it was fantastic and memorable to expand our knowledge of where these ants can be found, and it is always fun to have that rush of discovery when you find a new ant you haven’t seen before.

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

At the moment, I am teaching environmental sciences at the University of Alberta Augustana Campus. I work my research into the conservation parts of the course and try and engage students by exhibiting how there is still so much we can discover about the world around us.

In the past, I have done presentations with nature clubs to help spread knowledge about ants. A lot of people see ants as just annoyances in their garden or lawns, so teaching them that they are diverse, socially complex, and ecologically interesting is essential to helping them see ants as part of a wider world.

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

I think understanding the simple ecology and biology of social insects is still incredibly important. What are these insects feeding on? What are their effects on the ecosystem? What relationships do they have with other species? How do changes in the environment affect them? If we take the time to fully understand individual species, we can better understand their influence on the world and how the world influences them. This basic biological understanding becomes even more critical when considering the conservation of biodiversity and trying to prevent human-driven extinctions, as it gives necessary information to work from.

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

Tanya Huff’s Confederation novels. I would recommend them for science fiction readers, as they are a fun take on the “space marine” genre. It is a fun adventure series with good dialogue, exotic aliens, and an imaginative world.

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

I enjoy birding, especially in the winter when the ants are covered in snow. In Alberta, winter is when owls, such as Snowy or Great Greys can be found more easily, so it is a fun pastime in a land of white! I also play a lot of soccer and have been playing on the same team, with the same group of guys, since I was a teenager. Additionally, I enjoy having aquariums and terrariums, and at the moment have Imitator Dart Frogs (Ranitomeya imitator). Sadly, my ant-keeping skills could use some work.

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

Life is full of challenges; some can be fun, and some can be difficult. If I am in a particularly tough spot, I often take a break, reassess how to fix the issue, and then try to fix it. I have also been known to rant about the problem to my wife, who lovingly listens and helps me solve it. If things are really tough, it is always nice to get out and just watch nature, be it ants, birds, or other animals.

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

1. Gear to collect ants because there could be an interesting species! 2. A lot of clean, fresh water so that I don’t dehydrate. 3. A satellite phone so that when I am out of food/water and have found all the ants, I could call someone to come pick me up.

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

John Acorn. He is a professor at the University of Alberta. He was my master’s supervisor and is a great friend. He taught me to think critically, write succinctly, and how to observe the living world. His guidance and mentoring have benefitted me in all facets of my life.

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

Work hard towards your goals, but make sure you take time for other things in life. It is often easy to get bogged down in details, when there is so much more happening around you. Read every day. And lastly, seek out experts, ask questions, and don’t be afraid to actually ask those questions.

Interview with a social insect scientist: Graham Thompson

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IS: Who are you and what do you do?

Who am I? For the past ten years, I have been a Professor of Biology at Western University in Canada. My research programme folds-in a lot of social insect research, to the extent that we have dubbed our lab ‘The Social Biology Group’. This is a fun moniker for us that I hope reflects our interest in the ideas that permeate the study of social breeding systems, rather than any one person (me) or taxon. My lab usually supports between 3-5 people – though currently, we are at ten – and for the non-human taxa in our lab, we like to mix it up! We keep honey bees, termites and even fruit flies as playthings and models to test cool, evolutionarily-minded ideas in sociobiology.

 

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Our lab logo – The Social Biology Group at Western University (Canada). For the Study of Behavioural Genetics and Sociobiology.

 

IS: How did you end up researching social insects?

I got lucky. In my final year of undergrad, I was inches away from graduating into the void of nothingness. I took a field course that the University of Guelph (Canada) offered at the time, in Jamaica. The course was a bit of a junket, I guess, but did expose many of us to tropical ecosystems for the first time. Most students frolicked in the foreshore of Discovery Bay, but I was one of the few students involved in a non-SCUBA project. I had freshly taken a course with IUSSI member Prof. Gard Otis and, having been intrigued by social insects in class, I found the arboreal Nasutitermes nests in Jamaica to be just the thing! Fatefully, on that same field course was my future MSc supervisor, Prof. Paul Hebert (who some may recognize for his later work on DNA barcoding). We ran some impromptu allozyme gels (it was the 90s) to decipher each termite colony’s breeding genotypes. I think I was literally the first undergraduate to actually use a Punnet square in real life, or so it seemed. It was amazing! Chance meets opportunity. I got into grad school. I’ve been living the dream ever since.

 

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Talking termites at the Biology and Genomics of Social Insects Meeting in Cold Spring Harbour, May 2018, featuring IUSSI members Mackenzie Lovegrove, Ed Vargo, Anna Chernyshova and, in the background, Guy Bloch.

 

IS: What is your favorite social insect and why?

I don’t really have one, or, at least, not a permanent one. But I can remember the glee of first finding Glyptotermes in rotten logs shortly after moving to Melbourne (Australia), as well as Coptotermes in giant mounds along the roadside near Canberra, Drepanotermes hoarding dried grass in their underground chambers in the Outback, and massive Neotermes in a suburban wind-blown Eucalypt. I also felt centred living along-side the mighty Mastotermes darwiniensis on campus at James Cook University in Queensland (where I once did a postdoc). This termite is a fascinating creature from another time, and sadly, is on its way out after a 200 M year reign. Maybe just a few million years left before its phylogenetic branch falls off completely? But these and other fleeting moments of biophilia do not tempt me to rank any taxon above another. Did I even mention bees yet? Plus, if I promote my favourite taxon at the expense of yours, as some do, inadvertently, with informercial-style talks at conferences, then I run a risk of coming across as a taxonomic chauvinist. I, therefore, think it’s best to keep our field taxonomically (and otherwise) diverse.

 

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Seek and you shall find: ‘Termite Avenue’ near Cairns Australia with fellow termite enthusiasts (L-R) Vernard Lewis, Becky Rosengaus, and Susan Jones. This photo was taken from an impromptu excursion from IUSSI 2014. A farmer had ‘planted’ termite mounds along his driveway as if they were hedges. #onlyinAustralia

 

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

As the late Ross Crozier once said, science is a grand tapestry of which we are all threads. Or something along these lines…anyway, when my Ph.D. supervisor was once asked this same question, he, being somewhat more poetic than me, conveyed to the interviewer that for him, discovery was as much about the people he was funding and teaching and training as it was about the biological discoveries they were making. Like Ross, I also view science as a social enterprise and my best memories are not necessarily centred around making single discoveries – although I am indeed proud of a few things – but rather the shared experience of doing so with others who have traveled, however briefly, alongside my science journey. I have met many such people along the way!

 

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A conference lost in time. With former Hamilton Award winner, the late Ross Crozier, who was my Ph.D. supervisor (1997-2000), and past IUSSI President Ben Oldroyd, with whom I did a postdoc (actually, two! 2003-2007) and who taught me everything I know about honey bees. Ross and Ben have been great mentors to me.

 

IS: If teaching is part of your work, what courses do you teach?  Has your work on social insects helped to shape your teaching?

I teach third- and fourth-year courses at Western University in Canada. Currently, Animal Behaviour and Behavioural Ecology, which are both right up my alley. I do let loose on the details of honey bee biology from time to time, to the delight of my undergrads. But I only do so as one of many possible vehicles to teach the ways and means of natural selection. I strongly prefer to explain biological concept over any specific content, and usually emphasize process over memorized particulars.

 

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One of our apiaries at Western University in action! With students (L-R) Kyrillos Faragalla and Anna Chernyshova. I am ‘supervising’.

 

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

I recently read The Price of Altruism by Oren Harman (Norton; 2010) and am meandering my way through two books by philosopher and biographer Ullica Segerstrale: Defenders of the Truth (Oxford; 2000) and Nature’s Oracle (Oxford; 2013). I met Ullica at a recent conference, and she deserves great credit for chronicling the on-going history of our field. All three of these books are biographical narratives of the people – George Price, Bill Hamilton, Ed Wilson, and others – whose lives, luck, insight, and in some cases, misfortune, shaped the academic turf on which we now play. I love this stuff.

 

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The Price of Altruism is high, as Oren Harman explains.

 

IS: Did any one book have a significant influence in shaping your career?  What was the book and how did it affect you?

At risk of sounding unoriginal, it was, of course, The Selfish Gene! I read it as a senior undergraduate and, in a whiff, the whole of my undergraduate training made sense, albeit, retroactively. It is bemusing to think that biology degrees are still taught by rolling out information piecemeal, course by course, without a grand unifying theme that ties it all together. The Selfish Gene did that for me, as it famously has for so many others.

 

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Lab photo 2017 (L-R) Anna Chernyshova, me, Anthony Gallo, Kristin Ransome, Christine Scharf, Julia Saraceni, Rahul Choorakkat, Kyrillos Faragalla and, missing from this photo, our three beekeepers:  Rick Huismann, Andrew Pitek, and Alex Guoth.

 

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

Little known fact: I am an equestrian. I own my own horse and train with a coach to compete at shows. My horse and I are currently qualified for the Trillium Hunter Jumper Association Championships held in Toronto, the big end-of-season glitzy tourney for the top horse-rider pairs in the province (of Ontario). True story!

 

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Me and my trusty steed, a 16.3 hh Warmblood-cross gelding. On this day, he was Reserve Champion for his division.

 

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

I think the best quality for success in academia is sheer persistence. That, and playing the long game. Summer field season didn’t work out? -80C freezer went kaput? Paper sent to review for the third time? Funding pulled at the last minute? No problem! Dust yourself off and keep going as best you can. My advice, if you’re asking for it, is to stay consistent and persistent in your effort. We can all tolerate setbacks, even big ones, provided we recover quickly and keep going.

 

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One of our summer-long experiments in progress: feeding probiotic-infused pollen patties to increase performance and enhance resistance to disease. Or, we’ll see anyway!

 

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

Sewall Wright’s Evolution and the Genetics of Populations Vols. I-III. We’ve all read Vol. IV, so no need to bring that.

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

A mentor who infused me with confidence and direction? An inspiring colleague who made academia fun when, without them, it wouldn’t have been? An influential author who taught me all I need to know? A friend who rounded out your life away from the lab? Your lab mate who killed it and lifted everyone around them? For me, it’s all of these, all the time.

 

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Recruiting kindergarten beekeepers at Wortley Public School in London, Ontario.

 

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

Start early.

 

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It’s never too soon to begin your career in social insect research! My kids helping me tend the bees.

 

The Food Search Box assay – where do we go from here?

A blog post highlighting the article by N. Tsvetkov, B. Madani, L. Krimus, S. E. MacDonald, and A. Zayed in Insectes Sociaux

By Armo Zayed

 

Assay for spatial learning and memory

Assay for spatial learning and memory.

 

As central place foragers, honey bees have an amazing ability to fly several kilometers away from their colony to forage and then beeline back to their home without getting lost. Honeybee foragers can perceive and communicate spatial information via the famous waggle dance. But how are these traits encoded in the honey bee genome?

When Nadia Tsvetkov joined my lab in 2012, she was keenly interested in studying the genetics of spatial learning and memory in bees. She spent several months training bees to fly through mazes. The maze experiments were fun but took too long, and our sample sizes were far fewer than the hundreds of bees needed to tease out the likely subtle genetic effects on learning and memory. What we needed was an assay that was as fast and as easy to standardize as the proboscis extension reflex assay – the workhorse of insect olfactory learning and memory studies. We tried some different approaches (one involved a very beautiful but unwieldy maze constructed out of Christmas balls) until colleague Dr. Suzanne MacDonald, a vertebrate biologist at York University’s Department of Psychology, suggested that we try the food search task paradigm. The paradigm is commonly used to study spatial learning and memory in primates and rodents. A common protocol entails hiding toys or food in boxes within a testing arena that animals are allowed to explore. Over time, the animals learn and can recall the location of boxes that contain rewards.

So we set out to try a similar assay on bees. For prototyping, we used some common and inexpensive items; we made the testing arena of clear Tupperware containers and we employed several artificial flowers made out of Q-tips. After a few pilot assays, we decided that a small arena containing four flowers was the best compromise between complexity and length of the experiment. Nadia worked out the testing protocol that involved placing bees into the arena where one of the artificial flowers had a sucrose reward. Once a bee found and fed from the rewarding flower, it was removed and tested again for a total of three training trials. Finally, the bee entered the arena where none of the flowers had a sugar reward. This time, the bee had to rely solely on its memory to find the focal flower. Our data analysis showed that bees subjected to this test exhibited two telltale signs of learning and memory: they improved their ability to find the rewarding flower during training, and they were able to recall the location of the rewarding flower after training. The Food Search Box (FSB) was born.

We carried out two more experiments to test the utility of the FSB for studying spatial learning and memory in bees. We first compared the performance of nurses (young honeybee workers that nurse the brood) and foragers (older workers that forage outside the colony) in the FSB paradigm. While both nurses and foragers did equally well in the training trials, foragers did substantially better than nurses in the memory test. So, is it age (young vs. old) or behavioral state (nurse vs. forager) that is influencing spatial memory in the FSB? To answer this question, we carried out another study on same-aged workers. We treated these workers with either cGMP, which causes precocious foraging, or cAMP, which does not alter behavioral state. The cGMP-treated bees performed similarly to foragers in the FSB, while cAMP and the control bees performed like nurses in the assay. Taken together, the results of these two experiments indicate that behavioural state (nurse vs. foragers) is primarily associated with differences in spatial memory in the FSB.

We are very excited by the results of the FSB; we were able to test bees quickly without much attrition. It is feasible to screen hundreds of bees within a short period, opening up the door for genetic and genomic studies of spatial learning and memory in honeybees. While it is certainly possible to improve on the design to enhance automation (i.e., RFID readers or tactile sensors to passively record visits to artificial flowers), the low-tech version presented in the paper is very easy to set up and perform. We are looking forward to feedback from the community on the test, and we hope it will provide a useful tool for studying spatial learning and memory in honeybees and other insects.

Ants colonise bird nests and raise broods in them

A blog post highlighting the article by M. Maziarz, R. K. Broughton, G. Hebda, and T. Wesołowski in Insectes Sociaux

By Marta Maziarz

As an ornithologist, I have focused on the reproduction of birds but often overlooked the fact that bird nests can also be home to many invertebrates that find shelter, food or a suitable microclimate within them. When we discovered ant workers and their larvae inside nests of the wood warbler Phylloscopus sibilatrix, curiosity drove us to study this phenomenon.

An initial literature review revealed just a handful of published records of ant broods found inside bird nests, including blue tits Cyanistes caeruleus breeding in nest-boxes in Corsica (Lambrechts et al. 2008), and great tits Parus major and marsh tits Poecile palustris occupying tree cavities in primeval stands of the Białowieża Forest, Poland (Mitrus et al. 2015). Blem and Blem (1994) reported ant colonies on the side of nests in nest-boxes used by prothonotary warblers Protonotaria citre but gave no further details. This surprising scarcity of observations of ants in songbird nests suggested that this phenomenon may be exceptional and occur only among cavity-nesting species.

Our discovery of ant workers and their larvae in wood warbler nests, which are domed structures composed of dry grass, moss, and leaves and situated on the forest floor, challenged this view. We made the original finding during long-term studies of wood warbler ecology in 2004-2015 in Białowieża Forest (Eastern Poland), which prompted us to document this phenomenon systematically during 2016-2017. In 2017, we also contacted researchers in Switzerland and the UK to ask them to inspect nests for the presence of ants and their broods. We wanted to find the frequency of ants colonising wood warbler nests, and whether ants are present in wood warbler nests elsewhere in the species’ breeding range.

During our systematic observations in 2016-2017, we found adult ants in 43% of warbler nests, and one-third of nests also contained ant larvae or pupae. These ant broods were situated within the sidewalls of the nests, at or just above ground level. The most frequent species were Myrmica ruginodisor M. rubra, and occasionally Lasius niger, L. platythoraxor L. brunneus. These numbers, compared to 30% of nests containing adult ants and 20% containing broods during the earlier (2004-2015) period, indicated a long-term association between the ants and the birds. The findings from Białowieża Forest contrasted with those from Switzerland and the UK, where we only found single cases of adult ants and their broods. The different frequencies of ant presence between regions could be due to varying densities of bird or ant nests between woodlands transformed by humans to a different degree, but further studies would be necessary to confirm this.

These first records of adult ants and their broods in wood warbler nests showed that occupation of bird nests by ants can be a locally common phenomenon, which may have been overlooked previously in this and other songbirds. Systematic examination of nests belonging to different bird species would be valuable in understanding this further.

Furthermore, the occurrence of ant broods in the walls of wood warbler nests showed that ants colonised these structures following their construction by birds. Why they do this remains unclear; are the ants attracted to the nests by their structure, the presence of other invertebrates as a source of protein, or by heat generated by the birds? More work is underway to answer these questions, but it seems that these potential ant-bird interactions could be much more widespread than has been suspected.

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Wood warbler nests are dome-shaped and constructed of leaves, grass, and moss. They are usually hidden among low herb vegetation, under a tussock of grass or sedge, or wedged under fallen branches or logs. Such structure and locations could promote their occupation by ants, for example, Myrmicaspp., which raise their broods in similar places.

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Numerous ant Myrmicaspp. larvae and two larger, well-grown blowfly Protocalliphoraspp. larvae (centre-right) in the wall material of a wood warbler nest

References

Blem CR, Blem LB (1994) Composition and microclimate of Prothonotary warbler nests. Auk 111:197–200.

Lambrechts MM, Schatz B, Bourgault P (2008) Interactions between ants and breeding Paridae in two distinct Corsican oak habitats. Folia Zool 57:264–268.

Mitrus S, Hebda G, Wesołowski T (2015) Cohabitation of tree holes by ants and breeding birds in a temperate deciduous forest. Scand J For Res 31:135–139.

Microbiomes and worker tasks

Highlighting the article written by J. C. Jones et al. in Insectes Sociaux

Written by Insectes Sociaux Editor-in-Chief, Michael Breed

Molecular techniques for identifying microbial community composition have created a
true biological revolution. Recent discoveries lead us to understand the bacteria as an
evolutionarily complex and diverse domain, and this in turn has sparked interest in
characterizing microbiota from a large number of contexts. Of particular significance has been the exploration of gut microbiomes, which vary dramatically among species, and developmentally within species. Gut microbiomes interact strongly with diet and health, giving added interest to studies focusing on this subset of communities (Dunn 2011, DeSalle and Perkins 2016).

We have long understood the importance of the gut microbiome in social insect species. In termites, some components of the microbiota reduce cellulose to usable sugars while in other species, members of the microbiota fix nitrogen. More recent studies of ant and bee gut microbiomes have shown some level of intraspecific consistency even over broad geographic ranges, but also variation associated with diet and to a certain extent differences among colonies.

In this issue of Insectes Sociaux, Jones and her colleagues (Jones et al 2018) focus on
differences in the gut microbiota based on task group in honeybee (Apis mellifera) colonies. This is a question previously addressed by Kapheim et al (2015) but Jones and colleagues add critical dimensions by age-matching the worker bees in their study and collecting gut samples from bees observed performing specific tasks.

Each of five experimental colonies consisted of 1500 workers of the same age and from
the same source colony (400 of which Jones and colleagues individually marked). They
observed worker behavior in ten to fourteen-day old bees. Nurses, food receivers/handlers and foragers were noted and collected. This approach allowed assessment of diet and task-related differences in microbiomes independent of age-related developmental effects.

Jones et al (2018) found that Firm-4 (Lactobacillus mellis), one of the characteristic
bacteria of the honeybee microbiome, was more prevalent in nurse and food handling bees than in foragers. This pattern was also seen with quite a few other bacteria species, which had higher presences in nurses and/or food handlers than in foragers. One species, Lactobacillus kunkeei, was more common in forager guts, although they found it less commonly there, so this result is more provisional. Of particular note in the guts of food processing bees was Bartonella apis, as this species expresses genes that may be involved in the degradation of secondary plant metabolites.

Globally, the microbiomes of nurses and food handlers were more diverse than the
microbiome of foragers. Jones et al (2018) suggest that the needs for carbohydrate metabolism are higher for nurses and food handlers and that perhaps this drives functional differences in the gut microbiome between these task groups and foragers.

Concerns over bee health, responses of bees to diseases or parasites, and the impact on bees of the agricultural use of antimicrobials have generated much of attention given to bee microbiomes (Napflin and Schmid-Hempel 2018, Raymann and Moran 2018). While these topics are important, the microbiomes of social insects existed long before humans started to impact social species, and social insect microbiomes must have evolved alongside sociality. How might gut microbiomes facilitate worker task performance? Do they determine workers’ roles within colonies? The cause and effect relationship between task group and microbiome could go in either direction, with task environment driving the microbiota or the nature of the microbiological community feeding back into the task choice of bees. This study presents these alternatives as tantalizing avenues to pursue in future research.

References

DeSalle R, Perkins SL (2016) Welcome to the microbiome: Getting to know the trillions of bacteria and other microbes in, on, and around you. Yale University Press 264pp.

Dunn R (2011) The wild life of our bodies: Predators, parasites, and partners that shape who we are today. Harper 304pp

Jones JC, Fruciano C, Marchant J, Hildebrand F, Forslund S, Bork P, Engel P, Hughes WOH (2018) The gut microbiome is associated with behavioural task in honey bees. Insect Soc https://doi.org/10.1007/s00040-018-0624-9

Kapheim, KM, Rao VD, Yeoman CJ, Wilson BA, White BA, Goldenfeld N, Robinson GE (2015) Caste-specific differences in hindgut microbial communities of honey bees (Apis mellifera). PLoS ONE 10: e0123911

Napflin K, Schmid-Hempel P (2018) Host effects on microbiota community assembly. J Anim Ecol 87: 331-340

Raymann K, Moran NA (2018) The role of the gut microbiome in health and disease of adult honey bee workers. Current Opinion in Insect Science 26: 97-104