Nicholas W. Calderone Department of Entomology 6130 Comstock Hall Cornell University Ithaca, NY 14853 (607) 254-7417 (p) (607) 255-0939 (f) nwc4@cornell.edu

Field Lab:
Dyce Laboratory for Honey Bee Studies
203 Freese Rd.
(607) 275-0266

ACADEMIC RANK:
Associate Professor

APPOINTMENT:
75% Research – 25% Extension

EDUCATION:
Ph.D.   The Ohio State University - December 1988
M.S.    The Ohio State University - March 1985
B.S.     The Ohio State University - March 1977

AREAS OF SPECIALIZATION:
Evolution of social behavior and sociobiology
Honey-bee pests, parasites, and pathogens
Research Interests
Research Publicatons (Past 5 years)
Research Publicatons (submitted)
Extension Interests
CU Apiculture Extension Program
Extension Publications (Past 5 years)
Extramural Funding (Past 5 years)

RESEARCH INTERESTS:
Biology of honey bee pests, parasites, pathogens and predators: During the past decade, several new honey bee pests have come on the scene, and some old ones are presenting new challenges. One of the most challenging is the parasitic bee mite, Varroa jacobsoni, which threatens the continuation of commercial beekeeping and those sectors of the greater agricultural community that depend on honey bees for pollination.  Adult female mites are phoretic on adult bees, but reproduce exclusively on the immature stage of the bee. Mites enter the cells of immature bees just prior to their being capped. A mite’s reproductive success is greatly affected by its choice of both adult and immature hosts. Mites entering drone cells can expect to produce 2.6 offspring, while those entering worker cells can expect to produce 1.3 offspring. Those entering queen cells will not produce any offspring. Similarly, mites selecting a nurse bee as an adult host will be brought to the heart of the brood nest where new immature hosts are abundant. Those that find themselves on foragers are not brought into proximity with new immature hosts as often. In addition, because foragers have a brief lifespan, mites on foragers have a high risk of dying outside the nest.  Not surprisingly, host-seeking female mites exhibit considerable discrimination in their choice of both adult and immature hosts.  Females are found much more frequently on nurse bees than on foragers. In addition, they are found in drone cells more often than workers cells. They are almost never found in queen cells. During the past several years, my colleagues and I have been investigating the host-seeking behavior of the adult female mite. We have been seeking to identify the chemical and physical stimuli that the mite uses to find appropriate adult and immature hosts.  We have found that female mites can differentiate between nurse bees and foragers, and we are examining the behavioral repsonses of mites to nurse bee and forager kairomones. We are currently isolating the specific chemicals involved in each of these processes.

The organization and evolution of insect societies: Insect societies engage in numerous activities, including the construction, maintenance and defense of a nest; the location, collection and storage of food; and the rearing of offspring. Many species also reproduce by swarming. These activities are performed in a social context, involving extensive coordination among hundreds, thousands, or even millions of individuals, many performing the same task, others performing different tasks. In most cases, these tasks are interconnected; that is, the performance of one task either depends on, or affects the need for, the performance of another task. This dynamic connectivity presents a colony with significant organizational challenges. A colony must possess a coherent task structure to ensure the efficient and reliable performance of tasks. It must also possess mechanisms that ensure that workers are allocated among the various tasks in appropriate numbers. These challenges are complicated by the fact that the colony’s needs and the environment in which the colony lives are in a constant state of flux and by the fact that workers have limited neural processing capabilities.            

           The organizational structure of most social systems falls somewhere along a continuum between highly centralized and completely decentralized. Centralized control is a common feature of many human endeavors and typically involves a ‘management’ caste. Centralized control is rare among the social insects, being confined to species with small colony populations. These cases usually involve dominant-subordinate relationships between queens and workers.  If the queen is able to assess the status of the entire hive, she may direct, or stimulate, the activities of a small number of individual workers.  For example, queens of Lasioglossum zephyrum direct where returning pollen foragers should deposit their load. In the larger societies of the termites, honey bees and many species of wasps and ants, centralized control over the daily activities of workers would be impossible, impractical or inefficient. Previous models for the organization of work in insect societies have emphasized, to greater or lesser degrees, a worker’s age or the need of the colony, or, more recently, genotypic differences among workers. These models have identified important factors affecting division of labor. They have not, however, led to a detailed mapping of a colony’s task structure revealing how the activities of individuals are integrated into an efficient and reliable workforce. Nor have they provided an explanation for the ability of the colony’s workforce to respond in a timely and appropriate manner to rapidly changing environmental conditions, although they do explain some of the constraints on individual and colony flexibility.

           Understanding the organizational structure of a complex biological system is a long-term goal that requires concentrating on one subsystem at a time. After the different subsystems are understood, one can attempt the more difficult task of integrating them into a whole system – the colony.  Of particular interest in this process is the determination of exactly what properties must characterize the individual subsystem components – the workers – that enable the subsystem to function. Of equal interest is discovering whether or not subsystems can be integrated into a whole colony using only the same properties that apply to individual workers within a subsystem. The honey bee society provides an ideal organism with which to pursue the long-term goal of understanding the organization of social systems. The honey bee society possesses a highly complex social structure; yet, it is easily manipulated for experimental purposes. There is also significant work being done on different aspects of the honey bee society’s organizational structure and on physiological and genetical attributes of individual workers. This related research provides a context that will allow for a more rapid and empirically based understanding of the colony’s task structure.

           My current research examines one section of a colony’s task structure – pollen collection. This examination takes place at several levels. First, it explores how a colony integrates information about diverse stimuli that affect its need for pollen into an appropriate behavioral response. This is an essential first step because it will reveal which stimuli are integrated into decisions affecting resource acquisition, an essential component of colony fitness, and if/how these stimuli interact with each other in affecting that decision, a measure of system complexity. Second, it examines the individual-level activities that contribute to the colony-level decision-making process. Several options exist. Individual foragers may directly assess each of the stimuli affecting pollen collection, integrating that information into an individual decision as to whether or not to collect pollen or nectar. Or, non-foragers may assess the level of different stimuli affecting pollen collection, then transmit this information to foragers for synthesis. Or, foragers may combine information that they acquire directly with information they receive from non-foraging nestmates in making their foraging decisions. Third, it seeks to identify the specific stimuli assessed by each of the different players in the pollen collection system. Fourth, it examines how workers acquire information about the colony’s need for pollen and how they disseminate that information to nestmates.

Integrated Pest Management of honey bee pests, parasites, pathogens and predators: Over the past several years, I have been working on optimizing the use of formic acid for control of V. jacobsoni. Recently, I have developed a delivery method that results in 95% efficacy as a fall treatment in the northeast. I have also found that effective treatment with formic acid in the fall does not have deleterious effects on honey bee colonies. I am working to confirm these finding and examine the effect of environmental variability on the effectiveness of this delivery system. I am also working on the development of new solid polymer delivery systems for essential oils with acaricidal activity. I have also examined various methods for estimating mite levels in honey bee colonies as part of the sampling protocol necessary to implement treatment decisions. Both ‘ether rolls’ and passive mite collection devices provide reasonable estimates of mite levels. I am currently working on ways to incorporate these methods into a decision making process for beekeepers.

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RESEARCH PUBLICATIONS (PAST 5 YEARS):
     Calderone N. W. Effective fall treatment of Varroa jacobsoni (Acari: Varroidae) in colonies of the honey bee (Hymenoptera: Apidae) with formic acid in a northern climate (in press)
      Kuenen   L. P. S. and Calderone  N. W. (2000) Varroa mite infestations in elevated honey-bee brood cells: effects of context and caste. J. Ins. Beh.  (in press)
      Imdorf A., Bogdanov S.,  Ochoa R. I. and  Calderone N. W.  (1999) Use of Essential Oils for control of Varroa jacobsoni in honey bee colonies. Apidologie 30: 209-228.
      Calderone N. W. and Nasr M. (1999) Evaluation of a formic acid formulation for the fall control of Varroa jacobsoni (Acari: Varroidae) in colonies of the honey bee Apis mellifera (Hymenoptera: Apidae) in a temperate climate.  J. Econ. Entomol. 92: 526-533.
      Calderone N. W. (1999) Evaluation of formic acid and a thymol-based blend of natural products for the fall control of Varroa jacobsoni (Acari: Varroidae) in colonies of Apis mellifera(Hymenoptera: Apidae). J. Econ. Entomol. 92: 253-260.
      Calderone, N. W. (1998) Proximate mechanisms of age polyethism in the honey bee, Apis mellifera. Apidologie 29: 127-158.
      Calderone N. W. (1999) Evaluating subsampling methods for estimating numbers of Varroa jacobsoni mites (Acari: Varroidae) collected on sticky-boards. J. Econ. Entomol. 92: 1057-1061.
      Kuenen  L. P. S. and Calderone  N. W. (1998) Positive anemotaxis by Varroa mites: responses to bee odour plumes and single clean-air puffs. Physiol. Entomol. 23: 255-264
      Calderone  N. W. and Turcotte  R. (1998) Development of sampling methods for estimating infestation levels of Varroa jacobsoni (Acari: Varroidae) in colonies of the honey bee, Apis mellifera (Hymenoptera: Apidae) J. Econ. Entomol. 91: 851-863.
      Calderone, N. W., Wilson, W. T, and Spivak, M. S. (1997) Evaluation of plant extracts for control of the parasitic mites Varroa jacobsoni (Acari: Varroidae) and Acarapis woodi (Acari: Tarsonemidae) in colonies of Apis mellifera (Hymenoptera: Apidae). J. Econ. Entomol.90: 1060-1086
      Kuenen, L. P.S. and Calderone, N. W. (1997) Transfers of Varroa mites from newly emerged bees: preferences for age- and function-specific adult bees. J. Ins. Behav. 10: 213-228

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RESEARCH PUBLICATIONS (submitted):
      Vaughan D. M. and Calderone N. W. Direct assessment of colony need by honey bee foragers.
      Calderone N.W.<, Kuenen, L.P.S. & Lin, S. Effects of sex, comb-cell type, airborne volatiles and surface extreacts from worker and drone larvae and cocoons on infestation of Apis mellifera brood by Varroa  jacobsoni. & Kuenen, L.P.S. Differential tending behavior by nurse bees towards worker and drone larvae in the honey bee, Apis mellifera      
      Calderone N. W. and Lin S. Chemical and physical determinants of colony need by honey bees, Apis mellifera.
      Calderone N. W. & Lin S. Repellant effects of royal jelly extracts consistent with the differential invasion by Varroa jacobsoni in honey bee (Apis mellifera) worker and queen brood cells.
      Calderone N. W. & Johnson, B. A biphasic within-nest behavioral pattern by honey bee pollen foragers in pollen-rich and pollen-deprived colony environments.
      Lin S. & Calderone N. W.  Responses of Varroa jacobsoni mites to extracts of honey bee nurses and pollen foragers in choice and no-choice tests.

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CU APICULTURE EXTENSION PROGRAM:
The CU Apiculture Extension Program is designed to meet the needs of beekeepers, the larger agricultural community and the general public. These needs are addressed through a series of programs, the centerpiece of which is THE MASTER BEEKEEPER PROGRAM.  This program is designed to train people interested in acquiring the basics skills required to become successful hobbyist and commercial beekeepers.  The Master Beekeeper Program consists of three levels of training - apprentice, journeyman and master beekeeper. The Apprentice Level requires the completion of a spring and fall workshop series. The Journeyman Level requires completion of a core group of workshops and a number of elective workshops. The Master Beekeeper Level requires the completion of an additional set of core requirements and elective workshops. The workshop series provides thorough training in a diverse range of applied topics in apiculture. Certification at each level is based on receiving a passing grade in a comprehensive examination. People wishing to attend the worksjops, but not interested in obtaining certificaiton, are encouraged to attend. A current schedule for the Master Beekeeper workshop series can be found at the CU Apiculture Extension Home Page.

The Cornell University Apicluture Extension Program also provides training to other state organizations with an interest in Apiculture, fruit and vegetable growers who depend on honey bees for pollination or who apply pestcides to their crops, and to the general public. The program provides training for state apiary inspectors through the PPP IDENTIFICATION AND MANAGEMENT CERTIFICATION PROGRAM. This program trains and certifies beekeepers in the proper procedures for diagnosing and treating the major bee diseases. Wildlife Specialists with the the New York State Department of Enviropnmental Conservation receive training in HONEY BEES AND BEARS. This workshop is designed to provide edcuation on the problems bears can cause beekeepers in the state. Wildlife specialists will be instructed on the identification of bear damage and on methods that beekeepers can use to prevent bear damage before it occurs. DEC Pesticide Specialists receive training in HONEY BEES AND PESTICIDES and DOCUMENTING PESTICIDE DAMAGE TO HONEY BEES. These workshops provide education on the relationship between honey bees and pollination, and on the damage that honey bees can sustain as a direct consequence of the misuse of pesticides. Participants receive training on the proper use of pesticides around honey bees, and on management practices that growers and beekeepers can use to prevent or minimize pesticide damage to honey bees. Pesticide Specialists also receive instruciton on the proper protocol for documenting pesticide damage to honey bees and collecting samples for laboratory analysis. These workshops are available to NYS Certified Pesticide Applicators for re-certification credit.

Many fruit and vegetable growers throughout the state depend on honey bees for pollination. Many apply pesticides that can affect honey bee populaitons. The Cornell University Apicluture Extension Program offers an educaitonal workshop on HONEY BEES AND POLLINATION. This workshops provides information on the role of honey bees as pollinators, how to maximize the benefits from honey bees and how to care for them while they are on site. Fruit and vegetable growers are also encouraged to particpate in the HONEY BEES AND PESTICIDES workshop.

       The general public contacts stinging insects in a variety of ways. Form any, the primary interests is in avoiding these creatures. The Apiculture Extension Program provides information on theidentification and treatment of stinging insects, including honey bees, yellow jacketsd, hornets and wasps. Included in this area is informaiton on removing bees from buildings and handling swarms. A workshop on HONEY BEES AND SOCIETY is also available.

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EXTENSION PUBLICATIONS (PAST 3 YEARS):
      Calderone N. W. (2000) Master Beekeeper Program Manual: Apprentice Level, Spring and Fall Courses (500 pp).
      Morse R. A & Calderone N. W. (2000) The value of honey bee pollination the United States. Bee Culture 128: 1-15
      Calderone N. W. (2000) IPM - wax moths, mice, wasps & robber bees.  Bee Culture 128: 32-35.
      Calderone N. W. (1999) IPM for Parasitic Honey Bee Mites. Bee Culture 127:
      Calderone N. W. (1999) IPM: Minimize Stress – Promote Health. Bee Culture 127: 33-36
      Calderone N. W. (1999) An Introduction to Integrated Pest Management for Honey Bee Pests. Bee Culture 127: 27-30
      Calderone N. W. (1998) Identification and management of Varroa jacobsoni in the northeast (4 p Cornell Cooperative Extension Fact Sheet)
      Calderone N. W. (1998) Sampling protocols for honey bee parasites and pathogens (2 p Cornell Cooperative Extension Fact Sheet)

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EXTRAMURAL FUNDING (PAST 5 YEARS):
USDA-NRI-CGP. Physical and chemical determinants of Varroa mite invasion of honey bee brood cells. $126,000.00 awarded for 11/1/2000 - 10 - 31/2002
USDA - Northeast Region Sustainable Agriculture, Research, and Education Program (SARE). Integrating natural products and genetic resources for control of Varroa jacobsoni, a parasitic mite of the honey bee, Apis mellifera L. $82,412.00 awarded for 9/1/2000 - 8/30/2002
USDA-Northeast IPM Program. Integrating natural products and genetic resources for control of Varroa jacobsoni, a parasitic mite of the honey bee, Apis mellifera. $82,412.00 awarded for 10/1/2000 - 9/30/2002
Organic Farming Research Foundation. Natural products and non-chemical methods for controlling parasitic mites in honey bee colonies. $6,275.00 awarded for 1/2000 - 12/2001
USDA-NRI-CGP. Physical and chemical determinants of Varroa mite invasion of honey bee brood cells. $72,000.00 awarded for 10/1/98 - 9/30/2000
USDA-Northeast IPM Program. Project coordinator and co-principal investigator with Prof. L. S. Willett of Cornell University on research project: A reduced pesticide IPM strategy for control of the parasitic honey bee mite, Varroa jacobsoni. $100,000.00 awarded for 6/30/97 - 7/1/2000
USDA-NRI-CGP. Host-location by the parasitic honey bee mite, Varroa jacobsoni. $116,000.00 awarded for 10/1/96 - 9/30/98

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