Research Interests

h Biological invasions of aquatic communities: controls and impacts

h Plankton ecology, particularly in reservoirs and large rivers

h Biological assessment of water quality

The primary focus of my research is population and community ecology in freshwater environments. My students and I use surveys and experiments to explore the factors that limit the distribution and abundance of aquatic organisms. My primary study organisms have been the freshwater zooplankton, the small animals that eat algae and serve as food for young fish.  In recent years, I have focused much attention to dispersal and how that regulates community structure in lakes and ponds. Studies of aquatic invasive species provide a handy vehicle for this research.  Favorite study organisms include a tropical cladoceran (Daphnia lumholtzi) and the Chinese mystery snail (Cipangopaludina chinensis), both of which have been highly successful invading lakes and rivers of the coterminous United States.  A recent sabbatical provided me the opportunity to study these and other invasive species in the numerous lakes of northern Wisconsin.  Other collaborative research has explored planktonic communities of the Missouri, Mississippi, and Ohio rivers.  Finally, my students and I have an ongoing interest in assessing water quality of lakes and streams of the Missouri Ozarks.

My graduate students have allowed me to explore many more questions than I would have done alone. Some students have studied other aspects of species invasions, particularly the role of interactions with native species. Other grad students have examined water quality and impacts of global climate change on aquatic invertebrates. Each of my students writes a detailed research proposal, presents their research at professional meetings, and is encouraged to publish.  Most of the 19 students who have completed a Master’s thesis under my direction (listed below) have completed their thesis in about two years, many have received awards and grants, and all have jobs in the field of biology or environmental science.  This employment success rate attests to both their motivation and the employment prospects related to aquatic ecology.


Collaborative External Grants

Planktonic indicators of reference conditions in great rivers: analyses for the EPA EMAP (funded by Environmental Protection Agency, with numerous collaborators), 2004-08.

Spatial distribution and dynamics of algae blooms in Ozark reservoirs (funded by Missouri Department of Natural Resources, with R. Rhodes, MSU), 2001-02.

James River total maximum daily loads (funded by Missouri Department of Natural Resources, with R. Pavlowsky and R. Biagioni, MSU), 2001-02.

Vertical migration and niche partitioning: tower experiments (fellowship from Max Planck Institut für Limnologie, with W. Lampert, MPI), 1998-99.

Missouri River Post-flood Evaluation: connectivity of rivers and their floodplains (funded by Missouri Department of Conservation, with numerous collaborators), 1994-96.

Invasion of an exotic cladoceran into North America (funded by National Science Foundation, with J. O'Brien and V. Smith, University of Kansas), 1994-97. 

Graduate Students                       Prospective Graduate Students

                                                                 Click above link for more information

Current Graduate Students 

Elizabeth Glidewell (2011-present):  Adaptation of an invading cladoceran to temperature. 

Kari Reynolds (2010-present): Cyanobacteria dynamics and their relation to algal toxins and taste and odor compounds in lakes. 

Whitney Kelley (2010-present): Impacts of size-selective predation by rusty crayfish on invasive and native snails.

Jason Wolf (2006-2010):  Effects of fish cues and turbidity on zooplankton vertical migration behavior. (On leave, thesis in progress 2010)


Graduate Students Completing a Thesis (all Master of Science, Biology, Missouri State)

Sarah Davis (2008-2010):  Effects of toxic metals on growth and recognition of predator-cues in freshwater snails. Present position: Environmental Specialist, City of Springfield, Missouri. 

Rachel Heth (2008-2010):  Impacts of road-crossing disturbance on stream macroinvertebrate communities. Present position: Ph.D. student, University of Missouri.

 Garrett Clark (2007-2009):  Distribution, growth, and competitive impacts of the exotic Chinese mystery snail (Bellamya chinensis) in the James River, Missouri. Present position: completing HS teacher certification, Missouri State University.

Kelli Dickerson (2005-2008):  Zooplankton in flowing waters. Present position: Environmental Scientist, Leigh Environmental, Springfield.

Angela Bandy Coats (2005-2007):  Top-down control of algal biomass in streams: the role of snails in the Upper White River Basin, Missouri and Arkansas.  Present position: Ecologist, Arkansas Department of Environmental Quality, Little Rock.

Kim Medley (2002-2004):  Zooplankton species composition and diversity in floodplain ponds: effects of flooding and local site characteristicsPresent position: Ph.D. student (Conservation Biology), University of Central Florida..

Tina Tamme Hopper (2000-02): Environmental limits to invasion of ponds by Daphnia lumholtzi Sars. Present position: Laboratory Supervisor, Missouri State University.

Mark Penticuff (1999-2002):  Biomonitoring of stream water quality: comparative sensitivities of macroinvertebrate indices and leaf decomposition rates. Present position: Science Teacher, Springfield Public Schools.

Kristen Pattinson (1999-2001):  Blue-green algae and the seasonal succession of Daphnia. Present position: Environmental Specialist, Missouri Department of Natural Resources.

Lori Soeken Gittinger (1996-98): The effect of turbidity on the distribution and life history of river zooplankton. Present position: Research Specialist, Illinois Natural History Survey, Brighton, IL.

Jennifer Cacka Graham* (1996-98): Competition between exotic and native cladocerans. Present position: Research Scientist, US Geological Survey, Lawrence, Kansas. (*now Ph.D.)

Matt Eisenbacher (1995-98): Effects of the exotic cladoceran Daphnia lumholtzi (Sars) on the growth rate and prey selection of bluegill sunfish (Lepomis macrochirus Rafinesque). Present position: Environmental Scientist, Leigh Environmental, Inc., Springfield, MO.

Robin Hurtubise Calfee (1993-96): Effects of UV-B radiation on freshwater invertebrates. Present position: Research Scientist, US Geological Survey, Columbia, MO.

Jocelyn Korsch (1992-97): Biomonitoring a stream impacted by wastewater discharges using macroinvertebrate diversity and bioassays. Present position: Park Naturalist, Missouri Department of Natural Resources.

Robert Schulz (1991-96) The Central Stoneroller, Campostoma anomalum, as an indicator of heavy metal contamination, using otolith age and growth analysis. Present position: Fisheries Biologist, Missouri Department of Conservation.

Frederick Dry* (1990-92): Effects of odonate predation on a littoral prey community. Present position: Assistant Professor, Arkansas State University.  (*now Ph.D.)

Katherine Koontz Perkins (1990-92): Effects of landfill leachate on the behavior, feeding rate, and growth rate of the prosobranch snail Elimia potosiensis. Present position: Laboratory Coordinator, Ozarks Technical Community College.


Refereed Publications        

For a reprint, email

Bukaveckas, P.A., A. Aufdenkampe, J.H. Chick, J.E. Havel, R. Schultz, T.R. Angradi,  D.W. Bolgrien, T.M. Jicha, and D. Taylor.  2011. Phytoplankton abundance and their contribution to particulate organic matter in the Ohio, Upper Mississippi, and Missouri Rivers.  Aquatic Sciences, in press.

Havel, J.E. 2011. Survival of the exotic Chinese mystery snail (Cipangopaludina chinensis malleata) during air exposure and implications for overland dispersal by boats.  Hydrobiologia, in press. (DOI: 10.1007/s10750-010-0566-3)

Gray, E., S. Nunziata, J.W. Snodgrass, D.R. Ownby, and J.E. Havel. 2010. Predation on green frog eggs (Rana clamitans) by Ostracoda.  Copeia 2010: 452-456.

Chick, J.H., A.P. Levchuk, K.A. Medley, and J.E. Havel. 2010. Underestimation of rotifer abundance a much greater problem than previously appreciated.  Limnology and Oceanography Methods 8: 79-87.

Havel, J.E. 2009. Stanley Ivan Dodson: A distinguished ecologist, naturalist, and teacher. Hydrobiologia 636: 1-6 (DOI: 10.1007/s10750-009-9960-0).

Dickerson, K.D., K.A. Medley, and J.E. Havel. 2009. Spatial variation in zooplankton community structure is related to hydrologic flow units in the Missouri River, USA.  River Research and Applications 25: 1-14.

Havel, J.E., K.A. Medley, K.R. Dickerson, T.R.. Angradi, D.W. Bolgrien, P.A. Bukaveckas, and T.M. Jicha. 2009. Effect of main-stem dams on zooplankton communities in the Missouri River (USA).  Hydrobiologia 628: 121-135.

 Havel, J.E.  2009. Cladocera. Pages 611-622 in G. E. Likens, editor. Encyclopedia of Inland Waters. Elsevier, Oxford.

 Havel, J.E., and R.J. Rhodes. 2009. Spatial disconnection of plankton dynamics in an Ozark reservoir. Lake and Reservoir Management 25: 1-11.

 Soeken-Gittinger, L.A., J.A. Stoeckel, and J.E. Havel. 2009. Differing effects of suspended sediment on performance of native and exotic Daphnia. Freshwater Biology 54: 495-504.

Medley, K.A., and J.E. Havel.  2007. Hydrology and local environment influences on zooplankton communities in floodplain ponds.  Wetlands 27: 864-872..

Havel, J.E., and J. Graham.  2006. Complementary population dynamics of exotic and native Daphnia in North American reservoir communities. Archiv für Hydrobiologie 167: 245-264.

Havel, J.E., and W. Lampert.  2006. Habitat partitioning of native and exotic Daphnia in gradients of temperature and food: mesocosm experiments. Freshwater Biology 51: 487-498.

Havel, J.E., and K.A. Medley. 2006. Biological invasions across spatial scales: intercontinental, regional, and local dispersal of cladoceran zooplankton. Biological Invasions 8: 459-473.

 Havel, J.E., J.B. Shurin, and J.R. Jones.  2005. Environmental limits to a rapidly spreading exotic cladoceran.  EcoScience 12: 376-385.

 Havel, J.E., C. E. Lee, and J. Vander Zanden.  2005. Do reservoirs facilitate invasions into landscapes?  BioScience 55: 518-525.

Havel, J.E., and J.B. Shurin.  2004.  Mechanisms, effects, and scales of dispersal in freshwater zooplankton: a synthesis.  Limnology and Oceanography 49: 1229-1238.

Havel, J.E., and K.R. Pattinson.  2004.  Spatial distribution and seasonal dynamics of plankton in a terminal multiple-series reservoir.  Lake and Reservoir Management 20: 14-26.

Shurin, J.B., and J.E. Havel.  2003.  Hydrologic connections as dispersal routes for the spread of the exotic cladoceran Daphnia lumholtziBiological Invasions 4: 431-439.

 Pattinson, K.R., J.E. Havel, and R.G. Rhodes.  2003.  Invasibility of a reservoir to exotic Daphnia lumholtzi: Experimental assessment of diet selection and life history responses to Cyanobacteria. Freshwater Biology 48: 233-246.

Havel, J.E., J.B. Shurin, and J.R. Jones.  2002. Estimating dispersal from patterns of spread: Spatial and local control of invasion by Daphnia lumholtzi in Missouri lakes. Ecology 83: 3306-3318.

Johnson, J.L., and J.E. Havel. 2001. Competition between exotic and native Daphnia: In situ experiments. J. Plankton Res. 23: 373-387.

 Havel, J.E., and J. Stelzleni-Schwent. 2000. Zooplankton community structure: The role of dispersal. Verh. Internat. Verein. Limnol. 27: 3264-3268.

Havel, J.E., J.K. Colbourne, and P.D.N. Hebert. 2000. Reconstructing the history of intercontinental dispersal in Daphnia lumholtzi by the use of genetic markers. Limnology and Oceanography 45: 1414-1419. 

Shurin, J.B., J.E. Havel, M.A. Leibold, and B. Pinel-Alloul.  2000. Local and regional species richness: A scale-independent test for saturation. Ecology 81: 3062-3073.

Havel, J.E., E.M. Eisenbacher, and A.A. Black. 2000. Diversity of crustacean zooplankton in riparian wetlands: Colonization and egg banks. Aquatic Ecology 34: 63-76.

Galat, D.L., L. H. Frederickson, D. D. Humberg, K.J. Bataille, J.R. Bodie, J. Dohrenwend, G.T. Gelwicks, J. E. Havel, D.L. Helmers, J.B. Hooker, J.R. Jones, M. Knowlton, J. Kubisiak, J. Mazourek, A. McColpin, R.D. Semlitsch, and R.B. Renken. 1998. Natural and controlled flooding of channelized Missouri River wetlands: Complementary processes for restoring river-floodplain connectivity.  BioScience 48: 721-733.

 Hurtubise, R.D.,  E.E. Little, and J.E. Havel. 1998. Methods for assessing the impacts of UV-B radiation on aquatic invertebrates.  Pp. 31-44 in Little, E.E., A.J. DeLonay, and B.M. Greenburg (eds.) Environmental toxicology and risk assessment. Vol. 7.  ASTM STP 1333, American Society of Testing and Materials, Philadelphia.

Hurtubise, R.D., J.E. Havel, and E.E. Little. 1998. Effects of ultraviolet-B radiation on freshwater invertebrates: Experiments with a solar simulator.  Limnology and Oceanography 43: 1082-1088.

 Havel, J.E., M.C. Barnhart, and J. Schnake Greene. 1997. Experimental investigations of water quality: The bioassay. American Biology Teacher 59: 349-352.

Havel, J.E., W.R. Mabee, and J.R. Jones. 1995. Invasion of the exotic cladoceran Daphnia lumholtzi into North American reservoirs. Canadian Journal of Fisheries and Aquatic Sciences 52: 151-160.

 Havel, J.E., and B.L. Talbott. 1995. Life history characteristics of the freshwater ostracod Cyprinotus incongruens and their application to sediment toxicity tests. Ecotoxicology 4: 50-62.

 Chaplin, J.A., J.E. Havel, and P.D.N. Hebert. 1994. Sex and ostracodes. Trends in Ecology and Evolution 9: 435-439.

 Havel, J.E., and P.D.N. Hebert. 1993a. Clonal diversity in parthenogenetic ostracodes. Pages  353-368 in: McKenzie, K.G., and P.J. Jones (eds.). Ostracoda in the Earth and Life Sciences, Balkema Publishers, Rotterdam.

Havel, J.E., and P.D.N. Hebert. 1993b. Daphnia lumholtzi from North America: Another exotic zooplankter. Limnology and Oceanography 38: 1837-1841.

 Havel, J.E., J. Link, and J. Niedzwiecki. 1993. Selective predation by Lestes sp. (Odonata, Lestidae) on littoral microcrustacea. Freshwater Biology 29: 47-58.

 Havel, J.E., P.D.N. Hebert, and L.D. Delorme. 1990a. Genetics of sexual Ostracoda from a low arctic site. Journal of Evolutionary Biology 3: 65-84.

 Havel, J.E., P.D.N. Hebert, and L.D. Delorme. 1990b. Genotypic diversity of asexual Ostracoda from a low arctic site. Journal of Evolutionary Biology 3: 391-410.

 Havel, J.E., and P.D.N. Hebert. 1989. Apomictic parthenogenesis and genotypic diversity in Cypridopsis vidua (Ostracoda, Cyprididae). Heredity 62: 383-392.

 Havel, J.E., C.C. Wilson, and P.D.N. Hebert. 1989. Reproductive investment and sex ratios in the viviparous onychophoran Plicatoperipatus jamaicensis. Oikos 56: 224-234.

 Dodson, S.I., and J.E. Havel. 1988. Prey response in Daphnia: Growth and life history. Limnology and Oceanography 33: 1276-1287.

Havel, J.E. 1987. Predator-induced defenses: A review. Pages 263-278 in Kerfoot, W.C., and A. Sih. Predation: Direct and indirect effects on aquatic communities. University Press of New England, Hanover.

 Havel, J.E., and S.I. Dodson. 1987. Reproductive costs of Chaoborus-induced polymorphism in Daphnia pulex. Hydrobiologia 150: 273-281.

 Havel, J.E. 1985a. Predation of common invertebrate predators on long-and short-featured Daphnia retrocurva. Hydrobiologia 124: 141-149.

 Havel, J.E. 1985b. Cyclomorphosis of Daphnia pulex spined morphs. Limnology and Oceanography 30: 853-861.

 Havel, J.E., and S.I. Dodson. 1985. Environmental cues for cyclomorphosis in Daphnia retrocurva. Freshwater Biology 15: 469-478.

 Havel, J.E., and S.I. Dodson. 1984. Chaoborus predation on typical and spined morphs of Daphnia pulex: Behavioral observations. Limnology and Oceanography 29: 487-494.


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Copyright 1999 John E. Havel
This page last updated February 06, 2011.