Research Interests
I study evolution at the interface of behavioral ecology, ecology, and development of animals. I am particularly interested in learning how ecological factors influence the evolution and development of individual behavior and form, and in turn, how these individual characteristics influence population-wide characteristics. As a model system, I have chosen social insects. Their remarkably plastic responses to changing environments make them ideal subjects for investigating the role of developmental plasticity in the evolution of behavior and form. With social insects, I can study changes at three levels of organization: individuals, colonies, and populations. Responses that interest me include immediate changes in the behavior of the workers and queens and long-term changes in colony developmental characteristics, such as social structure and colony relatedness, size at maturity, life history trade-offs, and interactions with the local environment.
My current research focuses on a long-term developmental response of social insects, namely alternative colony-founding strategies. In many species, colonies produce large queens that are able to disperse, found new colonies and raise new workers on their own (independent-founding), while other species produce small queens that enter established colonies and produce offspring with the help of workers already present (dependent founding). Some species manifest alternative founding strategies, producing both kinds of queens, each with its attendant morphologies and behaviors. I found such a polymorphism in the cavity-nesting ant Temnothorax longispinosus. Analyses of fat content in virgin queens revealed three queen morphs: a small morph with almost no fat, a large morph with low fat, and a large morph with high fat. In contrast with the two larger morphs, small queens had little to no flight activity in mating swarms, which facilitates return to the natal nest after mating. I also found that when the mother queen is absent, the colony generally produced large new queens. Queen presence correlated with nest densities in different forest patches, revealing an important connection between ecological factors, and the physiology and development of individuals. Colonies from different populations varied in their response to queen presence. In high-density populations in which independent founding has low success rates, large queens were still produced when mother queens were removed, but fewer were produced and they had reduced independent-founding abilities. In this case, selection has not changed the response to queen absence, but has shifted other traits associated with that response.
Research on reproductive alternatives in social insects is having great impacts on our understanding of variation in social insect behavior and colony structure. Ants in which dependent founding is common or fixed tend to have dense populations of colonies lacking clearly defined colony boundaries, the ability to out-compete other ant species for resources, low dispersal and gene flow, and are more likely to be destructive invasive species. In contrast, independent-founding species have lower nest densities and greater dispersal between suitable habitats. Because the extent to which social insects dominate terrestrial ecosystems often depends on the founding strategy, understanding reproductive behaviors will yield valuable insights into ecosystem properties.
I have worked on several other projects studying the development and behavior of social insects. As an undergraduate, I found evidence of queen-worker conflict over colony sex ratios in two populations of the ant Temnothorax curvispinosus, in which the population with more queens per nest allocated more resources to male production. I have also studied foraging and prey capture behavior of army ants in the tropical rainforests of the Philippines, Kenya, and Panama while supported by a Watson Fellowship. For my masters research, my advisor, Robert Jeanne, and I studied trade-offs in colony foraging behavior in a social wasp in Costa Rica. Foraging rates were tightly correlated with larval numbers, which ultimately depended on the number of workers at nest initiation and the proportion of workers parasitized. I also found that colonies adjusted foraging rates as the colony aged, shifting workers from foraging to nest defense. There was a clear advantage of larger colony size: large colonies were able to raise their first workers faster and thereby increase the chances of raising new workers before interruption by predators. I also lead a side project with several researchers studying swarm recruitment behavior during emigrations by the paper wasp Apoica pallens in Costa Rica. Unlike most swarming wasps that mark a scent trail on substrates, A. pallens scouts have a unique calling posture that likely releases airborne pheromones, attracting the rest of the colony to a new nest site.
I will expand my research program in the future to include additional aspects of developmental alternatives in social insects. Among the questions I will be investigating are: 1) what are the communication pathways within colonies that relay environmental pressures to behavioral/developmental responses? 2) how do interactions with other species of ants and other animals affect colony responses? 3) when is plasticity adaptive? 4) what affects do parasites and symbionts have on allocation decisions? 5) how do alternative founding strategies influence gene flow and population subdivision? 6) what effect does each alternative founding strategy have on the surrounding ecological community? The results of these projects will provide a clearer picture of how and why ant communities and populations differ, and what factors contribute to specialization on one founding behavior, to population sub-division, and to differing colony organizations, all of which have profound effects on the interacting ecological community.
Student learning will be a major goal of these projects. One of the advantages of social insects is their versatility for answering a wide range of interesting biological questions. The cavity-nesting ant species that I study are easily collected, their upkeep in lab is extremely simple and successful, and their small individual and colony size makes housing large numbers possible in a small area and at low cost. Students can develop a large number of valuable skills working with social insects, through behavioral observation within and among colonies, physiological analyses of reproductive and non-reproductive individuals, community ecology studies on species interactions related to colony characteristics, and DNA analyses to study population characteristics (gene flow, genetic structure, speciation) or within colony characteristics (relatedness measures, intracellular parasites). Cavity-nesting ants can be found almost anywhere in the continental United States, making local projects possible and opening up appealing opportunities for science education outreach to the local community.Teaching Philosophy
"What we play is life" - Louis Armstrong.
When I read Louis Armstrong's quote about jazz music, it makes me think of one of my own great passions in life: teaching. The chance to inspire creative expression and a passion for knowledge in others, to have an outlet for my creativity and enthusiasms, and to shape our future by preparing students to be scientists and scholars motivates me to teach. My current teaching philosophy rests heavily on three important principles:
1) Teaching is an evolving process, in which a variety of methods are refined, enlivened, and sometimes discarded (Handelsman et al., 2004). Determining which methods are successful relies on the use of student-centered and goal-directed assessment (Wiggins, 1998). It also means that the teacher will never become boring or complacent. Just as testing a hypothesis usually raises more questions than it answers, trying out a new teaching method can open up a world of possibilities for improvement. As part of this process, I plan to try a number of different techniques ranging from lively lectures to individual problem solving to collaborative learning activities. Lectures will include diverse mediums for communicating information, including broad outlines, clear visuals, live demonstrations, and video clips. I also want to engage the students, framing the information in a way to which students can relate and find interesting, such as asking students to compare the peacock's tail to modern human adornments or ATP synthase to a water wheel powering a mill. Most students learn effectively by teaching and discussing a subject, so I plan to use group activities that encourage students to learn actively. This may be as simple as asking students to discuss a problem with a neighbor, or as complex as specially designed labs and field classes that allow students to apply knowledge to real biological questions. Since many biology classes do not have accompanying labs, I will encourage the application of science to real problems in the lecture class through investigative writing assignments and independent projects. I will also solicit feedback and constructive criticism from students at several points during the term, through the use of online anonymous surveys, evaluations, and discussions with students after completing an assignment. By assessing the effectiveness of a diversity of approaches, I will improve my teaching and learn what best stimulates and challenges students' creative abilities.
2) Enthusiasm generated in students and the freedom to explore are critical for stimulating a thirst for knowledge in students. I facilitate this by exuding enthusiasm for the subject I am teaching and encouraging students to find their own paths to learning. Once curiosity has been generated, students are more likely to motivate themselves to learn about the subject. In my classes, students will retain concepts by applying the terms, equations, or facts to biological mysteries that I introduce. Self-motivation encourages independent explorations, uncovering many new and related subjects. I first experienced the power of enthusiasm and exploration when I adapted a lab investigating the courtship behavior of live crickets to introduce animal behavior to introductory biology students. I began the lab by showing short video clips of spectacular courtship displays in a variety of other animals, discussing the important principles of communication and conflict. Then students set up observations on male crickets competing with each other, courting female crickets, and mating. The vicious competitions between males engaged students, and by the end of the lab they were cheering for their favorites. I was peppered with welcome questions about the behavior and many students stayed late to get extra observations completed.
3) Students are best prepared to be scientists by practicing science. Since I want to prepare students for future problems they will face, it is natural that they should learn concepts by applying them to independent investigations, both within classes and as part of mentored research. In the future I hope to teach general biology, entomology, behavioral ecology, and evolution courses within which students will learn science by practicing science, both in lectures and in labs. In particular, project-based labs will challenge students to investigate novel problems and discover their own solutions, while providing the real-world challenge of integrating results from their classmates' experiments (National Research Council, 2003). Using insects as model organisms can facilitate this approach. Insects are easy to keep in large numbers, have incredible diversity, and have short generation times. These characteristics allow students to generate interesting hypotheses and test them with sufficient sample sizes to interpret their own results. Mentoring students working on independent projects provides some of the best opportunities to give students valuable experience practicing science much as they would in future jobs and academic settings. My research on the evolution of development and behavior in social insect populations is sufficiently diverse to accommodate a wide variety of student interests from physiology and developmental biology to behavior and ecology. At the same time, the questions asked and the study techniques used are generally simple enough for students to understand in the often short time they will work with me. Some of my most rewarding experiences have come from collaborating with undergraduates and watching their growth as scientists as we design, implement, and discuss independent research projects. The best part of these experiences is when students work with me as a colleague, proposing novel ideas and giving unique perspectives that improve our research in addition to our learning experiences. My goal is for the students to come away from each class or mentoring experience better prepared to function as scientists and scholars.
In summary, my passion for teaching will stimulate new and innovative techniques to test in the classroom, and self-motivated and enthusiastic students becoming scientists. As a result, my students will be prepared to practice science in the future. Even though not all my students will become scientists as a profession, the ability to solve problems, think critically about mysteries in the natural world, and appreciate scientific progress will be vital to becoming a good citizen in today's world. This inspires me to teach.
Handelsman, J., Ebert-May, D., Beichner, R., Bruns, P., Chang, A., DeHaan, R., Gentile, J., Lauffer, S., Stewart, J., Tilghman, S.M., and Wood, W.B. (2004). Scientific teaching. Science 304(5670), 521–522.
National Research Council. (2003). BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: National Academies Press.
Wiggins, G.P. (1998). Educative assessment: Designing assessments to inform and improve student performance. San Fransisco: Jossey-Bass.
Curriculum Vitae
Mailing Address: Department of Entomology, University of Wisconsin ,
546 Russell Labs, 1630 Linden Dr. , Madison , WI 53706
E-mail Address: howard@entomology.wisc.edu
Work Phone: (608) 262-7136
Home Phone: (608) 255-0312
- Ph.D., expected May, 2007, Zoology, University of Wisconsin, Madison, WI
Thesis title: Alternative colony-founding behavior in the cavity-nesting ant, Temnothorax longispinosus
Committee: Robert Jeanne (advisor), Jeffrey Baylis, Janette Boughman, Anthony Ives, and Ken Raffa. - MS, 2003, Zoology, University of Wisconsin-Madison, Madison, WI .
Masters thesis title: Foraging and growth strategies of the swarm-founding wasp, Polybia occidentalis. - BS, 1998, High Honors in Biology, Davidson College, Davidson, NC .
Honors thesis title: Queen-worker conflict in Leptothorax ants.
- Teaching Assistant for Animal Biology Lab, Department of Zoology, University of Wisconsin.
January – May 2000, September 2004 - present - Teaching Assistant for Introduction to Biology, Department of Zoology, University of Wisconsin.
January – May 2004 - Benthic Aquatic Macroinvertebrate Technician, Duke Power Environmental Labs, Mecklenburg County, NC.
October 1997- June 1998 - Research Assistant, for Dr. Valerie Banschbach, Davidson College.
June 1996- August 1996
Teaching awards
- Wm. C. Burns Teaching Award, Department of Zoology, U. of Wisconsin, Madison, 2006
- Howard Hughes Medical Institute Teaching Fellowship, 2004, to participate in the Howard
- Hughes Medical Institute New Generation Program for Scientific Teaching and mentor an undergraduate student
Graduate research awards
- National Science Foundation Graduate Research Fellowship, 2000-2003. Effects of colony stage on worker tempo in swarm-founding wasps.
- National Science Foundation Graduate Research Fellowship Travel Grant, 2001. Foraging behavior during the pre-emergence period of the swarm-founding wasp, Polybia occidentalis
- Kinney Award, spring 2006. Travel to the International Union for the Study of Social Insects international meeting in Washington , D.C., July, 2006. Department of Entomology, University of Wisconsin
- Emlen Award, 2003. Queen dimorphism in Leptothorax longispinosus. Department of Zoology, University of Wisconsin
- John Jefferson Davis Research Award, 2001. Foraging behavior during the pre-emergence period of the swarm-founding wasp, Polybia occidentalis. Department of Zoology, University of Wisconsin
- University Graduate Fellowship, 1999-2000. University of Wisconsin-Madison
- Watson Fellowship, 1998-99. Recruitment to prey in Eciton, Dorylus, and Aenictus army ants. Thomas J. Watson Foundation
- Phi Beta Kappa, 1998
- Sigma Xi Excellence in Undergraduate Research Award, 1998
- Barry M. Goldwater Scholarship, 1997-98. For excellence in mathematics, science, and engineering
- Howard Hughes Summer Research Grant, summer 1997. Queen-worker conflict in Leptothorax ants. Davidson College
- Grants-in-Aid of Research, 1997. Queen-worker conflict in Leptothorax ants. Society of the Sigma Xi
- Howard, K.J. and D. Kennedy*(submitted). Mating behaviors of the queen polymorphic ant Temnothorax longispinosus. Ethology.
- Bouwma, A.M., K.J. Howard, and R.L. Jeanne. (in press) Rates of non-legionary ant predation on colonies of a swarm-founding social wasp. Biotropica.
- Howard, K.J. 2006. Three queen morphs with alternative nest-founding behaviors in the ant, Temnothorax longispinosus. Insectes Sociaux 53: 480-488.
- Howard, K.J. and R.L. Jeanne. 2005. Shifting foraging strategies in colonies of the social wasp Polybia occidentalis (Hymenoptera, Vespidae). Behavioral Ecology and Sociobiology 57: 481-9.
- Bouwma, A.M., K.J. Howard, and R.L. Jeanne. 2005. Parasitism in a social wasp: effect of gregarines on foraging behavior, colony productivity, and adult mortality. Behavioral Ecology and Sociobiology 59: 222-233.
- Howard, K.J. and R.L. Jeanne. 2004. Rates of brood development in a social wasp: effects of colony size and parasite infection. Insectes Sociaux 51: 179-185.
- Howard, K.J. , A.R. Smith, S. O'Donnell, and R.L. Jeanne. 2002. Novel method of swarm emigration by the epiponine wasp, Apoica pallens (Hymenoptera Vespidae). Ethology, Ecology, and Evolution 14: 365-371.
- International Union for the Study of Social Insects - International meeting - (invited paper). Queen presence determines female alate size in the queen-size polymorphic ant Temnothoraxlongispinosus. K.J. Howard. July 30, 2006
- International Union for the Study of Social Insects - North American Branch Meeting (submitted paper). Relationship of fat content, propagule numbers, and queen size to nest density in Temnothorax longispinosus. K.J. Howard. October 8, 2004
- Entomological Society of America National Meeting (submitted paper). Size dimorphism in queens of Leptothorax longispinosus. K.J. Howard. October 28, 2003
- Behavior Interdisciplinary Association, University of Wisconsin , Madison (invited paper). Wasps plan ahead: size of nest built by Polybia occidentalis swarms predicts colony foraging rates and productivity. K.J. Howard and R. L. Jeanne. Feb. 14, 2002
- Entomological Society of America National Meeting (submitted paper). Wasps plan ahead: size of nest built by Polybia occidentalis predicts pre-emergence foraging and productivity. K.J. Howard and R.L. Jeanne. Dec. 11, 2001
- National Conference for Undergraduate Research (submitted paper). Queen number and reproductive differences between Leptothorax curvispinosus populations at two southeastern sites. K.J. Howard, V.L. Banschbach, and P.A. Peroni. May 7, 1998
- North Carolina Academy of Science Annual Meeting (submitted paper). A comparison of queen number and reproductive allocation in two southeastern populations of the ant Leptothorax curvispinosus. K.J. Howard, V.L. Banschbach, and P.A. Peroni. April 20, 1998
- Entomological Society of America National Meeting (submitted paper). Queen number and reproductive differences between Leptothorax curvispinosus populations at two southeastern sites. K.J. Howard, V.L. Banschbach, and P.A. Peroni. Dec. 17, 1997
- Tri-Beta
- Entomological Society of America
- International Union for the Study of Social Insects
Guest lectures
- "Insect Defenses", Insect Behavior, Department of Entomology, University of Wisconsin February, 2005
- " Darwin 's Revolutionary Idea: Introduction to Evolution and Natural Selection". Introductory Biology, Department of Zoology, University of Wisconsin. April, 2004
Teaching Assistantships
- Animal Biology, led discussions and labs, Department of Zoology, University of Wisconsin. January-May 2000, September 2004-present
- Introductory Biology, led discussions and labs, Department of Zoology, University of Wisconsin. January-May 2004
Teaching and mentoring classes
- Awarded an HHMI Teaching Fellowship, 2004. As part of this fellowship, I participated in the Howard Hughes Medical Institute New Generation Program for Scientific Teaching. I also mentored 1 student working in the 2004 Symbiosis Summer Program organized by the Department of Plant Pathology, University of Wisconsin , while participating in the class Teaching Biology: Special Topics - Mentoring. June-August, 2004
- Inquiry-based Biology Teaching with instructors Jo Handelsman and Sarah Lauffer. Focused on scientific teaching, inquiry-based techniques, classroom dynamics and diversity, course design, and assessment. Fall, 2004.
- Delta Instructional Materials Development Seminar, January - May, 2004. Collaborated with Robert L. Jeanne to develop questions and assessment materials for the use of "clickers" (a.k.a. personal response systems) in the Introductory Biology lecture class. The class included collaborative learning about teaching as a scientific process, assessment, and diversity.
- Teaching College Biology, with instructor Wayne Becker. June 2002. Introductions to important teaching concepts including active learning, engaging students, syllabus preparation, lecture designs. Also included a variety of guest speakers from teaching and research institutions.
Mentoring experience
- Mentored 5 students working on summer research projects for independent research in the departments of Entomology and Zoology, U. of Wisconsin, May-August 2003, and 1 student May-August 2005. Students wrote and revised a final scientific paper analyzing their results and discussing their findings.
- Continued mentoring 1 student as she presented a poster entitled "Relationship of worker size to queen size in the queen dimorphic ant Leptothorax longipsinosus" as part of Introductory Biology Mentored Research Poster Presentation session, University of Wisconsin, December 10, 2003.
- Insect Ambassadors - Department of Entomology, U. of Wisconsin, Madison. Jan. 2005 - present
Dr. Robert Jeanne
University of Wisconsin-Madison, Department of Entomology
1630 Linden Dr., Madison, WI 53706
Phone: 608-262-0899;
Email: jeanne@entomology.wisc.edu
Dr. Jeffrey Baylis
8735 Hidden Valley Road, Cross Plains, WI 53528
Phone: 608-798-3602;
Email: jrbaylis@tds.net
Dr. Jo Handelsman
University of Wisconsin-Madison, Department of Plant Pathology
1630 Linden Dr.,
Madison, WI 53706
Phone: (608) 263-8783;
Email: joh@plantpath.wisc.edu
Dr. Janette Boughman
University of Wisconsin-Madison, Department of Zoology
428 Birge Hall,
Madison, WI 53706
Phone: 608-262-2636;
Email: jboughman@wisc.edu
Dr. Gale Oakes
University of Wisconsin-Madison, Department of Zoology
243 Noland Hall,
Madison, WI 53706
Phone: 608-265-5867;
Email: goakes@wisc.edu
