The Honey Bee Laboratory

at William Paterson University

Dr. David C. Gilley

Associate Professor of Biology

Why Study Honey Bees?

Bees are a model research organism. Honey bees have excellent memory abilities, flexible aging physiology, an unusual genetic system, extreme altruism, and symbolic communication abilities, making them a fascinating subject for study from molecular, physiological, ecological, and evolutionary points-of-view.  In my lab we investigate these unusual biological traits and their impact on the social organization of honey bee colonies.

Bees are important for agriculture. Approximately one-third of our food relies on honey bees for pollination.  Threats to the well-being of America’s honey bee colonies started well before Colony Collapse Disorder appeared in 2006.  These threats include mites, microscopic pathogens, pesticides, and the economic viability of the practice of beekeeping itself.  Colony Collapse Disorder compounds all these threats and continues to make research on honey bee biology an issue of national priority.  In my lab we study aspects of honey bee biology that may provide tools for more effective management of honey bees in an agricultural setting.


Honey Bee Foraging Pheromones

A successful honey bee forager performs a “waggle dance” upon returning to the hive to communicate to other bees the location of the food source she has discovered. The waggle dance is one of the few examples of symbolic language in non-human animals (the dance codes distance to the food source as the duration of the straight portion of the dance, and direction to the food source by the angle of the straight portion of the dance relative to the gravity vector.)  Waggle-dancing bees emit a blend of compounds inside the hive which appear to function like a foraging pheromone to stimulate a colony's foraging. To work out how these dance compounds function we use a combination of field and laboratory approaches.

Field studies allow us to investigate how this pheromone functions in a natural context. Here I supplement pheromone levels to provoke a foraging response by pipetting dance compounds into a dish to be inserted into the hive.

By marking bees for individual identification, we can track their responses to the pheromone, such as increasing their foraging flights to the feeder shown here

We use laboratory behavioral assays, such as this T-maze, to measure bees' responses to the dance compounds

Our initial discovery of this pheromone used gas chromatography and mass spectrometry to identify the compounds emanating from dancing bees

Honey Bee Foraging Behavior in an Urbanized Environment

Pollinators are in decline globally, one cause for which is urbanization of habitats that previously supported diverse and abundant pollinator communities. We are interested in how honey bees, as generalist pollinators that have evolved impressive abilities to discover and efficiently harvest floral food resources, cope with the urbanized environment surrounding New York City. Using our rooftop apiary and indoor observation hives, we are learning what plant species honey bees are foraging by trapping pollen from returning foragers, where honey bees are foraging by decoding their waggle dances, and how healthy these colonies are by keeping records of hive weight and status. Seasonal records of this information will allow us to investigate how the environment shapes plant-insect interactions, including mismatches between plant bloom and pollinator behavior as a result of climate change.

Waggle dances can be video recorded from observation hives and used to determine foraging locations. Other aspects of foraging behavior can also be studied.

Students Nicole Miller and Trevor Courtright inspect colonies on our rooftop apiary

Our observation hives, pictured here with an insulated cover, have access to the environment via tubes through the walls


Research Fields

Animal behavior

Honey bee biology

Kinship and cooperation


Foraging behavior

Research Goals

My research passion is working out the behavioral, physiological, and genetic mechanics underlying complex social interactions. Social insect colonies are particularly interesting subjects for this research because of their altruism, decentralized control, unusual genetic structures, complex communication among a large population of heterogeneous individuals. Honey bees, which have been the focus of my research for two decades, have the additional practical benefits of being easy to procure and maintain, as well as an economic rationale for the study of their behavior.

Professional History

2011 - present        Associate Professor of Biology, William Paterson University

2007 - 2011            Assistant Professor of Biology, William Paterson University

2003 - 2007            Research Associate, USDA Agricultural Research Service, Tucson AZ

2003                       Ph.D. Cornell University, Department of Neurobiology and Behavior

1997                       B.A.  Cornell University, Biological Sciences, magna cum laude

Selected Publications

  • Gilley, DC & Thom C (2015) Waggle-dancing bees emit body odors: new evidence of an important role for scent in bee foraging       communication. Bee Culture Sept 2015, 33-37.
  • Gilley DC (2014) Hydrocarbons emitted by waggle-dancing honey bees increase forager recruitment by stimulating dancing.  PLoS ONE 9(8):e105671.
  • Gilley DC, Kuzora JM, Thom C (2012) Hydrocarbons emitted by waggle-dancing honey bees stimulate colony foraging activity by causing foragers to exploit known food sources.  Apidologie 43:85-94.
  • Kronforst MR, Gilley DC, Strassmann JE, Queller DC (2008)  DNA methylation is widespread across social Hymenoptera. Current Biology 18(7): R287-288.
  • Thom C, Gilley DC, Hooper J, Esch HE (2007)  A scent to the waggle dance. PLoS Biol 5(9): e228. doi:10.1371/journal.pbio.0050228.
  • Gilley DC, Degrandi-Hoffman G, and Hooper J (2006) Volatile compounds produced by live honey bee queens. Journal of Insect Physiology 52(5): 520-527.
  • Tarpy DR, Gilley DC, and Seeley TD  (2004)  Levels of selection during queen replacement in honey bees: conflict, cooperation, or both? Behavioral Ecology and Sociobiology 55: 513-523.
  • Gilley DC  (2003)  Absence of nepotism in the aggressive interactions between workers and dueling queen honey bees.  Proceedings of the Royal Society B 270 (1528): 2045-2049.
  • Gilley DC  (2001)  The behavior of honey bees (Apis mellifera ligustica) during queen duels.  Ethology 107: 1-22.


Courses Taught: upper-level courses for majors


Animal Behavior (BIO 3500)

This course is a survey of the discipline of animal behavior. Both proximate and ultimate approaches to understanding behavior are discussed, along with the methodologies used to study behavior at both of these levels of analysis. Topics include the neural basis of behavior, the role of genes in behavior, learning and cognition, communication, sexual selection, parent-offspring conflict, competition, social behavior, altruism and kin selection.  This course prepares students for the advanced study of behavior and physiology while reinforcing core concepts of biology with which students are already familiar.


Zoology (BIO 3180)

This course is a study of the diversity of animal life from marine invertebrates to terrestrial arthropods, and includes the five major vertebrate groups: fishes, amphibians, reptiles mammals and birds. Special emphasis is placed on biological principles relevant to the modern study of zoology such as adaptation and natural selection, structure-function relationships, and developemental genetics. Students learn to identify members of major animal taxa as well as the relationships among these animal groups, and acquire skills in zoological laboratory techniques through field collecting, dissection, and live-organism research projects.


Parasitology (BIO 3990)

This course introduces students to the biology of parasitic organisms and the interactions of parasites with their hosts.  In addition to a survey of parasitic taxa (including single-celled eukaryotes, “worms”, and arthropods), topics include host defenses, evasion and manipulation of host defenses by parasites, pathogenesis, epidemiology, virulence, and parasite ecology.  Texts and lectures emphasize both proximate (physiological) and ultimate (evolutionary) perspectives on these topics.  This course prepares students for graduate study of invertebrate zoology, medical or veterinary parasitology, and epidemiology.


Evolutionary Medicine (BIO 4430)

Evolutionary medicine is an integrative discipline that applies evolutionary knowledge to the understanding of human biology, both normal and abnormal, to create a holistic understanding of how health and disease emerge. This course explores the roles of natural selection, genetic variation, developmental plasticity, and cultural evolution on human health traits such as immunity, metabolism, puberty, senescence, and psychology. Intended primarily for students considering medical and health professions, this course is designed to implement the recommendations for scientific foundations for future physicians made by the American Association of Medical Colleges, the Howard Hughes Medical Institute, and the National Academy of Sciences.


Biology Capstone Seminar (BIO 4800)

Restricted to senior Biology majors. The course requires each student to do an in-depth study of a selected topic to demonstrate their mastery of departmental learning objectives. This study requires gaining familiarity with a body of research literature, oral presentation of research literature, and preparation and revision of a professional-grade review paper. Additional assignments evaluate students’ scientific skills, assess their knowledge across the breadth of Biology, and prepare them for careers in Biology. This course is Writing Intensive.

Courses Taught: introductory and intermediate courses


Evolution, Ecology, Behavior (BIO 2490)

Introduction to the study of ecology, evolution, and behavior. Diversity and classification, history of life, evolutionary theory, population growth and regulation, species diversity and community structure, energy flow and nutrient cycling, ethology, behavior genetics, evolution of behavior, sociobiology, behavioral ecology. Required of biology majors.




General Biology 2 (BIO 1640)

The course includes principles of whole organism structure and function with emphasis on organ systems. Required of

biology majors.




Introduction to Animal Behavior (BIO 1400)

This course is an introduction to the scientific study of animal behavior. Students learn about the nature of science, the diversity of animal behavior, its evolution, its underlying physiological mechanisms, and the theoretical basis of our modern understanding of behavior. The course emphasizes the natural behavior of non-human animals but also considers the implications of the principles of animal behavior for humans and domestic animals. Labs focus on the scientific method, which students will practice by means of observational and manipulative experiments involving live animals in both laboratory and field settings. Not open to biology majors.


Human Biology (BIO 1200)

This is a course in biological science and research methodology from the perspective of the human body and its systems. The course lays the foundation of knowledge of biology: the chemistry of living organisms, the cell tissues, organs, organ systems and the organism; homeostasis as it applies to human survival; evolutionary processes; genetic, reproductive and other areas of biotechnology; and the human body systems and how they function and malfunction. Biomedical research, the use of humans as research subjects, and other bioethics issues are addressed. Research methodology includes use of the scientific method in laboratory exercises, and critical analyses of research studies. Laboratories include varied exercises in anatomy, physiology, genetics, and evolution. Not open to biology majors.

Courses Taught: graduate


Seminar in Organismal Biology (BIO 7700)

An advanced seminar designed to enable students to investigate recent advances in organismic biology.

Parasitology (BIO 5990)

This course is cross-listed with BIO 3990 (see above).

Evolutionary Medicine (BIO 5430)

This course is cross-listed with BIO 4430 (see above).


I chose to teach at a student-focused institution for the opportunity to mentor students in research.

Students in the bee lab typically participate in an existing project for a season, often working as part of a student team, and then may move on to independent research projects. They are involved in every stage of the scientific process including reading of background literature, experimental design, data collection, data analysis, and sometimes the publication process.

Interested in volunteering in my lab? See me or send an email.

Interested in independent research in my lab? See notes below.

Current Students


Trevor Courtright

Honors independent study project:

A Phenological Study of Honeybee Colonies and their Key Pollen-source Plants in the Urbanized New Jersey Landscape


Nikki Miller

Honors independent study project:

Investigating the foraging patterns of Apis mellifera in a mixed environmental setting

Bee Lab Alumni

Kristina Kupryk

Murat Mamkegh

Chad Socha

Kara Malone

Krystle Frederick

Jacqueline Kuzora

Corey Stein

Brian Smith

Ashley Miranda

Roy Feliciano

Erin Crehan

Raza Elias

Keith Malinak

Jessika Santamaria

Melissa Gallo

Alanna Nolan

Kevin Penkoski

Nina Ramdedovich

Virginia Hislop

William Manzo

Adalyne Arafet

Information for students considering independent research projects


Equipment and resources available

• On-site housing of live honey bee colonies in indoor observation hives (SciWest)

• On-site rooftop apiary of approx. 5 full-sized honey bee colonies (SciWest)

• Off-site apiary/fieldsite of approx. 5 full-sized bee colonies (Haledon)

• Equipment for lab studies of bees (e.g. incubator, cages, scopes)

• Equipment for field studies of bees (e.g., canopy, hive hardware, tables, feeders)


Examples of past student independent study projects

• What are the behaviorally active components of the honey bee waggle-dance pheromone?

• Does urbanization affect moth diversity in Northern New Jersey?

• Does the common pesticide imidacloprid affect honey bee colony health or productivity?

• Do honey bee swarms produce waggle-dance pheromone?

• How does the honey bee waggle-dance pheromone affect individual bee behavior?


Project considerations

• Projects may or may not involve honey bees but should utilize my expertise in animal behavior

• Recommended coursework before pursuing a project under my mentorship is BIO 2490 EEB and BIO 3500 Animal Behavior. Best time to express interest is during or after completing EEB, which should be well before your last year of undergraduate coursework. Other recommended upper-level courses are BIO 3180: Zoology, or BIO 3330: Field Entomology

• Most behavior projects do not require extensive training, but one semester in the bee lab as a volunteer or paid research assistant is recommended. Additional experience, such as a summer internship in the bee lab, is ideal

• Projects with bees (and many other animals in natural settings) will need to be completed over the summer or in the early fall because these animals are not active during the winter. Proposals for spring IS work need to consider these seasonal constraints.

• Students with known allergies to bee stings are not encouraged to work in the bee lab

• Note that an implicit goal of an Honor’s project in my lab is the production of a manuscript suitable for publication in an appropriate peer-reviewed journal such as American Journal of Undergraduate Research


Link to BIO 4990: Independent Study guidelines



Dr. David C. Gilley

Department of Biology

William Paterson University of New Jersey

300 Pompton Road

Wayne, NJ 07470


Office: Science Hall East 4038

Lab: Science Hall West 307

Phone: 973-720-2549

Email: gilleyd(at)wpunj(dot)edu


BIO WPU, September 15th 2014