Boudinot, who studied with major professor Phil Ward of the UC Davis Department of Entomology and Nematology, is the second UC Davis-affiliated entomologist to receive the honor in its 28-year history. Jessica Gillung of the Lynn Kimsey lab, Bohart Museum of Entomology, won the award in 2019.
Snodgrass (1875-1962), a leader in insect morphology, is known for his 76 scientific articles and six books, including Insects, Their Ways and Means of Living (1930) and the book considered to be his crowning achievement, the Principles of Insect Morphology (1935).
Boudinot has them all. “Principles of Insect Morphology and the Anatomy of the Honey Bee were the foundation of my studies,” he said. “I have both, plus his Textbook of Arthropod Morphology and Insects, Their Ways and Means of Living on my desk in the lab.”
The Snodgrass Award, which includes a certificate and cash prize, recognizes outstanding research by a PhD student who has completed a research thesis or dissertation in arthropod morphology, systematics, taxonomy, or evolution. Nominees are scored on honors, awards, achievements and recognition; recommendations of professors and advisors; grantsmanship, publications, creativity and innovation of thesis or dissertation; and contribution to morphology.
Boudinot's dissertation: “Systematic and Evolutionary Morphology: Case Studies on Formicidae, Mesozoic Aculeata, and Hexapodan Genitalia.”
He earlier received the prestigious Alexander von Humboldt Research Fellowship to do research on evolutionary and comparative anatomy in Jena, Germany. He will locate to Germany in early 2021 for the two-year fellowship, after completing intensive language studies.
'I Am a Morphologist Because of Robert Evans Snodgrass'
“I am a morphologist because of Robert Evans Snodgrass,” Boudinot wrote in his application. “Although I have had pressure from my earliest undergraduate to become a molecular systematist, it was my chance encounter with Snodgrass's Principles of Insect Morphology late one night in the college library that set the course of my career. I had struggled for years at that point to understand the biodiversity of insects and to untangle the deep morass of arcane terminology, but my vision was illuminated by the conceptual clarity, linguistic simplicity, and exceptional engravings of the Principles. This work continues to hold special dominion over my thinking, as it is through the principle of mechanical function for explaining comparative anatomical observations that I have come to my present understanding of systematic and evolutionary morphology.”
Boudinot wrote that his “career objective, in brief, is to resolve the morphological evolution of insects through the synthesis traditional morphology, as represented by Snodgrass, with recent trends in digital anatomy and bioinformatics. I envision a future wherein students rely not on Borror & Delong, a holdover from Comstock's 19thCentury manual, but rather learn about insect structure, function, classification, and evolution through manipulation of three- and four-dimensional digital anatomical models, substantiated via manual curation and dissection. I want students to see for themselves what I have understood through the study of Snodgrass's work, balanced by contemporary research: The origin of the Arthropoda and morphological transformation through geological time to the resplendent, and endangered, diversity of today.
“In sum, my identity as an entomologist, and as a scientist more broadly, is due to the insights on the language and phenomenology of morphological evolution I gained from the oeuvre of Snodgrass. Without these works, I would probably still be a botanist.”
Boudinot's research interests include the origin and evolution of complex phenotypic systems from the perspective of phylogenetic systematics, including molecular and paleontological evidence. Specializing on morphological identity and evolutionary transformation, the skeletomuscular system of Arthropoda, with emphasis on the male genitalia of Hexapoda and systematics of the Hymenoptera, particularly the Formicidae.
John Henry Comstock Award
Highly honored for his academics, leadership, public service activities, professional activities and publications while at UC Davis, Boudinot received the 2019 John Henry Comstock Award, the top graduate student award given by ESA's Pacific Branch. The branch encompasses 11 Western states, U.S. territories, and parts of Canada and Mexico.
In the Comstock award application, Steve Nadler, professor and chair of the Department of Entomology and Nematology, described Boudinot as “A highly respected scientist, teacher and leader with a keen intellect, unbridled enthusiasm, and an incredible penchant for public service.” Boudinot maintained a 4.00 grade point average and published 18 peer-reviewed publications on insect systematics, some landmark or groundbreaking work.
His most recent publications: one on Cretaceous Strepsiptera in Cladistics and the other on the iron maiden ants in Myrmecological News ("Two New Iron Maiden Ants from Burmese Amber (Hymenoptera:
Boudinot received multiple “President's Prize” awards for his research presentations at national ESA meetings. He organized the ESA symposium, “Evolutionary and Phylogenetic Morphology,” at the 2018 meeting in Vancouver, B.C. , and delivered a presentation on “Male Ants: Past, Present and Prospects” at the 2016 International Congress of Entomology meeting in Orlando, Fla.
Boudinot served on—and anchored—three of the UC Davis Linnaean Games teams that won national or international ESA championships. The Linnaean Games, now known as the Entomology Games, are a lively question-and-answer, college bowl-style competitions on entomological facts played between university-sponsored student teams.
Brendon served as president of the UC Davis Entomology Graduate Student Association from 2006 to 2019, and co-chaired the department's UC Davis Picnic Day celebration (with forensic entomologist Robert Kimsey) for three years.
Before enrolling in graduate school program at UC Davis in 2014, Brendon received his bachelor's degree in entomology in 2012 from The Evergreen State College, Olympia, Wash. Professor John T. Longino served as his mentor.
The paper appears in the current edition of Journal of Hydrobiologia.
“The water hyacinth, Eichhornia crassipes, isconsidered the world's most economically damaging aquatic weed,” said Bick, now a postdoctoral scholar at the University of Copenhagen, Denmark.
The free-floating perennial, native to the Amazon region of South America, is highly invasive throughout the world. It forms large floating mats when its roots and leaves intertwine. The aquatic weed is a major issue in the Sacramento-San Joaquin Delta in central California.
“This paper is the result of my first dissertation chapter,” said Bick, an agricultural entomologist who received both her master's degree (2017) and doctorate in entomology (2019) from UC Davis. “We aimed to determine if salinity was the reason N. bruchi was not effective at regulating the weed in the Sacramento-San Joaquin Delta compared with other worldwide locations. The results were not as clear cut as we hoped, as the study was limited in testing only adult weevils. However, the paper makes the case for including salinity as a screening variable for new biological control agents that are candidates for release in the Delta and other partially saline areas. Also, given the thoroughness of the experiments, there is at least one cool modeling paper to come out as a follow-up.”
The paper, titled “Effects of Salinity and Nutrients on Water Hyacinth and its Biological Control Agent, Neochetina bruchi, “was truly an all-hands-on-deck effort,” Bick said. "Specifically, a major project hurdle was the temperature in Davis."
She related that the greenhouse experiments on water hyacinth “weren't producing consistent results due to the high variation—and high heat--in water temperature.” So fellow scientists Danny Klittich, then a UC Davis doctoral student in entomology with the Michael Parrella laboratory, and Bob Starnes, then UC Davis senior superintendent of agriculture, built a giant water bath out of a leftover evaporative cooler from the Parrella lab.
Klittich is now the California Central Coast Agronomist with Redox Chemicals and chief executive officer and founder at HowToGrowRoses.org. Starnes is vice president of agriculture for UAV-IQ (Unmanned Aerial Vehicle Intelligence.
In addition to Klittich and Starnes, other co-authors are UC Davis postdoctoral scholar Elvira deLange of the Christian Nansen lab; then doctoral student Cindy Kron of the Frank Zalom laboratory; and undergraduate students Jessie Liu and Derrick Nguyen. Kron, now with UC Cooperative Extension, is the North Coast Area Integrated Pest Management (IPM) Advisor, serving Sonoma, Napa, Mendocino and Lake counties.
“Water hyacinth, Eichhornia crassipes (Mart.) Solms (Commelinales: Pontederiaceae), is an important aquatic weed worldwide. Previous studies demonstrate that releases of Neochetina bruchi Hustache (Coleoptera: Curculionidae) provide biological control in many locations, but not all. Notably, N. bruchi were unsuccessful at regulating water hyacinth in tidal brackish waters. Abiotic factors, including salinity and nutrients, affect water hyacinth growth, but little is known about the impact of salinity on weevil survival. We hypothesized that N. bruchi has a relatively low salinity tolerance. In a mesocosm experiment, we assessed weed growth in response to a range of salinity and nutrient concentrations. In a laboratory, we assessed adult N. bruchi mortality in response to various salinity concentrations. Results indicate that increasing nutrient concentration increases weed growth. When both nutrient and salinity levels were varied, nutrients increased leaf count, but not biomass, while salinity reduced growth and increased mortality. Increasing salinity concentrations increased adult weevil mortality; required concentrations were higher than that for weeds. Thus, these results did not provide support for the suggested hypothesis. Potential effects of salinity via other exposures to weevils need to be investigated. Elucidating abiotic factors important for weed growth and weevil survival may increase effectiveness of water hyacinth management practices.”
The water hyacinth was introduced in California in 1904. Scientists trace its history in the United States back to 1884 at the New Orleans Exposition. “Samples are said to have been given to fair-goers, and within 4 years, coastal fresh waters were infested from Texas to Alabama. By 1972, the infestation in Florida was estimated to be 200,000 acres,” according to Cornell University. “Large, floating mats of waterhyacinth obstruct navigation, clog irrigation works, disrupt the natural ecology of wetlands in many ways, exacerbate mosquito problems, and are costly to the tourism and recreation industries.”
Two biocontrol agents, weevils N. eichhorniae and N. bruchi, natives of Argentina, and surrounding areas in South America, were released in 1972 and 1974, respectively.
The question is crucial “because it is the foundation of essentially all biological questions,” says spider systematics researcher Lacie Newton, a doctoral student in the Jason Bond laboratory, UC Davis Department of Entomology and Nematology, and the lead author of newly published research that explores that question.
“For example,” Newton says, “making successful conservation efforts depends on knowing how to identify the threatened/endangered species from other closely related species that are not threatened.”
Her research on folding-door spiders or the Antrodiaetus unicolor species complex led to a journal article published in Molecular Ecology: “Integrative Species Delimitation Reveals Cryptic Diversity in the Southern Appalachian Antrodiaetus unicolor (Araneae: Antrodiaetidae) Species Complex.” UC Davis co-authors are Professor Bond, who is the Evert and Marion Schlinger Endowed Chair in Insect Systematics, and project scientist James Starrett of the Bond lab.
Folding-door spiders are so named because they close the entrances to their silk-lined burrows by pulling in the rim. They are often described as having stocky brown bodies, thick legs and large fangs.
The five-member research team, also including Professor Brent Hendrixson of Millsaps College, Jackson, Miss., and postdoctoral fellow Shahan Derkarabetian of Harvard University, used an integrative approach with several lines of evidence (morphological, behavioral, molecular, and ecological data) to form a consensus “about where we should draw the lines between species in this complex,” Newton said.
They targeted the Antrodiaetus unicolor species complex, which Newton said, are “great organisms for exploring species boundaries because even though these spiders do not have any obvious visual differences to tell them apart--with the exception of the smaller and lighter brown A. microunicolor-- there are significant genetic differences between certain populations, that is potential 'cryptic' species.”
Said Professor Bond of the journal article: “I think its significance lies in the innovative and multipronged approach (integrative) she employed to evaluating species boundaries. The study emphasizes the importance of using both genomic scale and ecological data rather than relying on traditional morphological features alone to delimit species. Understanding species boundaries is an imperative for cataloging and describing the planet's rapidly disappearing biodiversity.”
Newton won a second-place award for her oral presentation on species delimitation at the 2019 American Arachnological Society (AAS), held at Washington and Lee University, Lexington, Va. Her abstract: “Although species delimitation can be highly contentious, the development of reliable methods to accurately ascertain species boundaries is a fundamental and necessary step in cataloguing and describing Earth's quickly disappearing biodiversity. Species delimitation in spider taxa has historically been based on morphological characters; however, certain mygalomorphs are morphologically indistinguishable from each other yet have considerable molecular divergence." She is active in both AAS and the Society of Systematic Biologists.
First-Generation College Student
Newton, a first-generation college student, is a fifth-year doctoral program student whose research interests include systematics, Araneae, mygalomorph spiders, speciation pattern and process, phylogeography, molecular phylogenetics, and character evolution. She is the recipient of a year-long UC Davis Graduate Research Mentorship Fellowship that supports promising doctoral students that meet diversity criteria.
Born and raised in Eupora, Miss.-- “a very small town with less than 2000 people”--Lacie recalls a childhood that included “a significant amount of time outdoors with my family surrounded by the rich flora and fauna of the Coastal Plain Floristic Province.”
“This experience,” she related, “fostered my interest in biodiversity and later guided me to take additional science classes to learn more about the complexities of the living world.”
What sparked her interest in spiders? “I actually used to be terrified of spiders,” Lacie acknowledged. “It wasn't until fall semester of my sophomore year when I took a zoology course that I began to appreciate not only the vast amount of diversity within spiders but also how amazing they are as a group, such as the tensile strength of spider silk being comparable to steel, spider venoms playing a role in potential medical applications, and a myriad of feeding strategies, etc..”
“As my professor Dr. Brent Hendrixson shared his research interests (systematics of mygalomorph spiders and scorpions) and passion for scientific outreach, I evolved from a guarded student to a fascinated one. Additional summer field courses focused on the biology, evolution, and ecology of arachnids completely changed my career trajectory from becoming a medical doctor to an evolutionary biology professor with research emphasizing evolutionary processes of arachnid study systems, specifically mygalomorph spiders like Antrodiaetus.”
Career as Evolutionary Biologist
The UC Davis doctoral student plans a career as an evolutionary biologist, exploring the evolutionary history of mygalomorph spiders. “My ultimate career goal is to become a biology professor where I can perform research and teach in a vibrant academic setting,” Newton said. “As a professor, my aims are to become an expert in my desired field of evolutionary biology, continue to be involved in the scientific community through collaborations with researchers, and become an advocate for vision and change in science education. “
“To clarify,” she added, “I want to study the evolutionary history of arachnids by using emerging technologies/methods and bioinformatics tools. I also plan to participate in the scientific community by publishing articles in respected journals, attending and presenting at conferences, and collaborating with various researchers. Lastly, I plan to take part in teaching and outreach opportunities to convey my enthusiasm for science to others. I feel outreach is especially important to get children enthusiastic about science and to demystify science for the public.”
Newton aims to become a faculty mentor “who can positively impact students--the way my own undergraduate mentor Dr. Hendrixson affected my life--by using my position as a professor to extend opportunities to mentor high school students, undergraduate students, and graduate students, especially from underrepresented groups such as women and members of the LGBTQ community. Specifically, I want to mentor students about career options available as well as offer my own point-of-view about pursuing a career in a STEM field.”
At UC Davis, Newton served as a teaching assistant for the “Introduction to Biology: Biodiversity and the Tree of Life” course. She is mentor to undergraduate students in the Mentoring Program, Equity in Science, Technology, Engineering, Math, and Entrepreneurship (ESTEME) organization, a graduate student organization dedicated to improving equity and inclusion in STEM fields, entrepreneurship, and leadership positions. She also volunteers on the admissions committee for GOALS, the Girls' Outdoor Adventure in Leadership and Science, a summer science program for high school students to learn science hands-on while backpacking through the wilderness.
The "eight-legged wonders," as she said, fascinate her. It's "not only the vast amount of diversity within spiders but also how amazing they are as a group."
As environmental artists, bees are "responsible for the brilliantly colored flowers in our landscapes," and as environmental engineers, they engineer “the niches of multitudes of plants, animals and microbes.”
Page, with UC Davis roots and Arizona State University wings, has just authored a 256-page book, “The Art of the Bee: Shaping the Environment from Landscapes to Societies” (Oxford University Press), to be published Aug. 6.
“It's a long time in the making,” said Page, who received his doctorate in entomology at UC Davis and served as a professor and chair of the Department of Entomology (now Entomology and Nematology) before heading to Arizona State University (ASU), where he advanced to school director, college dean and university provost.
“Twenty-five years ago, my friend and mentor Harry Laidlaw (for whom the UC Davis bee facility is named) wanted to write a honey bee biology textbook,” Page recalled. When they finished the outline, “it looked very much like the excellent book by Mark Winston The Biology of the Honey Bee, published in 1987 by Harvard University Press. I decided we didn't need another one, and we still don't.”
The book differs in that it's a collection of “sparkling essays” that “read like mystery stories,” said Rudiger Wehner, professor and director emeritus of the Institute of Zoology, University of Zürich. “With these lucidly written stories, Page takes us on a delightful journey through the many biological traits that on the whole constitute the honeybees' social contract.”
“But don't be fooled by the amiable and personal style—the book is comprehensive—from colony collapse disorder to colony-level evolution—and chock full of the latest results, presented with clarity and depth, leavened with razor-sharp insights into social evolution,” noted Gene Robinson, director, Carl R. Woese Institute for Genomic Biology and Department of Entomology, University of Illinois at Urbana-Champaign.
Page said his book is geared toward “the person who has a basic knowledge of biology and a fascination with bees, perhaps an educated hobby beekeeper--there are a lot of them--or an undergraduate or graduate student with an interest.”
In addition to chapters on environmental artists and environmental engineering, Page includes chapters on social contracts, superorganisms, reproductive competitions, and concludes with “The song of the queen.”
In the epilogue, Page ponders the complexity of individual bees and their colonies, comparing them to humans. “Members of complex societies live close together in closed nests, shared home sites, villages, etc., or in closely connected nomadic tribes. As groups, they typically have a set of tacit rules by which they live that involves working for the good of the group, systems of group and resource defense, internal mechanisms of policing cheaters that don't cooperate and live by the rules, a division of labor often associated with group defense and gathering and sharing resources, and usually asymmetries and rules associated with reproduction. These same general characteristics seem to apply broadly across eusocial insects (aphids, termites, bees, ants, and wasps), eusocial rodents (naked mole rats), higher apes, and humans. Why? The similarities are inescapable due to the nature of social contracts; they must have specific elements to protect the power and will of individuals, whether citizens of the United States of America or workers in a honey bee colony. The contract binds individuals to a society, but the specific social organization evolves by reverse engineering. Natural selection acts on the whole colony; social structure evolves to fit the needs of the group within a given environment. “
Page points out that “Anthropocentric thinking can obscure the way we view nature and lead to false conclusions. Look at Aristotle and honey bee division of labor: For more than 2,000 years it was thought that the bees that work in the nest were postpubescent old men because they're hairy! In fact, the older bees forage and aren't hairy because the hairs break off as they age. I now see my work in a new light; we aren't so different, bees and humans. The elements of our social structures, and how they come about, have many similarities.”
Page is known for his research on honey bee behavior and population genetics, particularly the evolution of complex social behavior. One of his most salient contributions to science was to construct the first genomic map of the honey bee, which sparked a variety of pioneering contributions not only to insect biology but to genetics at large.
At UC Davis, he maintained a honey bee-breeding program for 24 years, from 1989 to 2015, managed by bee breeder-geneticist Kim Fondrk at the Harry H. Laidlaw Jr. Honey Bee Research Facility. They discovered a link between social behavior and maternal traits in bees.
UC Davis named him the 2019 distinguished emeritus professor. Nominator Steve Nadler, professor and chair of the UC Davis Department of Entomology and Nematology, praised Page as “a pioneer researcher in the field of behavioral genetics, an internationally recognized scholar, a highly respected author, a talented and innovative administrator, and a skilled teacher responsible for mentoring many of today's top bee scientists…he is arguably the most influential honey bee biologist of the past 30 years.”
Page has authored more than 250 research papers, including five books. Among them “The Spirit of the Hive: The Mechanisms of Social Evolution” (Harvard University Press, 2013) and “Queen Rearing and Bee Breeding,” with Harry H. Laidlaw (Wicwas Press, 1997). He is a highly cited author on such topics as Africanized bees, genetics and evolution of social organization, sex determination, and division of labor in insect societies.
Page, who received his doctorate in entomology from UC Davis in 1980, joined the UC Davis faculty in 1989 and left as emeritus chair of the Department of Entomology in 2004 when ASU recruited him for what would become a series of top-level administrative roles. He advanced from director of the School of Life Sciences to dean of Life Sciences; vice provost and dean of the College of Liberal Arts and Sciences; and university provost. Today he holds the titles of provost emeritus of ASU and Regents professor emeritus, as well as UC Davis department chair emeritus, professor emeritus, and UC Davis distinguished emeritus professor.
Page is an elected member of the American Academy of Arts and Sciences, the Brazilian Academy of Science, Leopoldina (the German National Academic of Science), and the California Academy of Science. He is a recipient of the Alexander von Humboldt Senior Scientist Award (Humboldt Prize, 1995), the Carl Friedrich von Siemens Fellowship (2013), James W. Creasman Award of Excellence at ASU (2018).
(Editor's Note: Geoffrey Attardo, assistant professor, UC Davis Department of Entomology and Nematology, published this piece July 29, 2020 on The Conversation website. This article is republished from The Conversation under a Creative Commons license. Read the original article.)
Bloodthirsty tsetse flies nurse their young, one live birth at a time – understanding this unusual strategy could help fight the disease they spread
Tsetse flies are bloodthirsty. Natives of sub-Saharan Africa, tsetse flies can transmit the microbe Trypanosoma when they take a blood meal. That's the protozoan that causes African sleeping sickness in people; without treatment, it's fatal, and millions of people are at risk due to the bite of a tsetse fly.
My entomology research focuses on insects that feed on the blood of people and animals. From a human health standpoint, understanding what makes all these bugs tick is key to developing ways to control them and prevent transmission of the diseases they carry, such as malaria, dengue, Lyme disease, West Nile virus and many others.
Tsetse flies stand out from their blood-feeding cousins the mosquitoes and ticks because of their unique reproductive biology. They give birth to live young and, even more unusual, the mother lactates and provides milk for her offspring. Here's how it all works – and why their unusual reproduction strategy might be a key to controlling tsetse flies and the parasite they carry once and for all.
From egg to larva
Scientists know of other flies that hold onto their eggs in their reproductive tract until they hatch into young larvae, with each brood consisting of dozens of offspring. The mother then tries to find a suitable source of nutrition in the environment, deposits the larvae and leaves them to survive on their own. The mother does not provide any nutrition for her young.
That's the standard fly way of life. Tsetse flies take a different approach.
Female tsetse flies develop just one single egg at a time. When the egg is complete, the mother moves it from her ovaries into her uterus in a process called ovulation. Once in the uterus, the egg is fertilized with sperm the female has stored in an organ called the spermatheca. While females can mate multiple times, they obtain all the sperm they need for their lifetime from a male fly during a single mating event.
After fertilization, the female keeps the egg in her uterus for five days while an embryo develops within the egg. When the embryo is ready, the egg hatches in the uterus of the female and the tsetse fly larva begins its life living inside its mother's uterus.
Milk meals for baby
Here's where tsetse flies dramatically diverge from most other insects.
Attached to the mother's uterus is a specialized gland that makes a milk-like substance. The organ is called the milk gland, and it produces a rich mixture of fats and particular proteins that provide the larva with all the nutrition it needs to develop into an adult.
Just like in mammals, the milk also transfers beneficial bacteria from the mother to the offspring. These bacteria are essential for tsetse flies, and without them adult female flies are unable to reproduce.
After five or six days of developing and feeding on milk, the larva is fully grown and ready to enter the world. The mother finds a safe spot and gives birth. The larva immediately burrows underground to avoid predators and parasites.
Once buried, the outer surface of the larva's skin hardens and turns black, forming a protective shell. This is called the pupal stage and it lasts for around three weeks. During this time, the pupa transforms into an adult fly.
It then emerges from the pupa, climbs out of the ground, and begins its life as an adult tsetse fly looking for hosts to blood-feed on and other tsetse flies to mate with.
Why live birth?
Why would an insect evolve this slow and resource-intensive way to reproduce?
One idea is that this method provides a defensive advantage relative to free-living larvae against parasites and predation. Larvae on their own have few (if any) ways to defend against these threats. But keeping larvae in the mother's uterus provides shelter and a guaranteed food source. While this strategy is much slower, scientists think the extra maternal care results in higher larval survival rates. It's a matter of quality over quantity.
A result of this reproductive strategy is that tsetse fly populations are small and slow to recover from control efforts, relative to more prolific insects like mosquitoes.
My colleagues and I hope that we can parlay our understanding of the molecular processes that regulate tsetses' milk production and mating behavior into new environmentally friendly, cost-effective and tsetse-specific control strategies for these insects.
The sleeping sickness tsetse flies spread is a potential issue for millions of people in 36 sub-Saharan countries, though the number of annual cases has decreased drastically thanks to major control efforts – including trapping flies, applying insecticides and releasing sterile males to the environment where they mate with wild females but don't produce offspring. Ultimately, we'd like to contribute to the World Health Organization's goal of eliminating African sleeping sickness by 2030 with a new way to prevent the transmission of disease-causing trypanosomes to people and animals./h2>/h2>/h2>/figcaption>/h1>