Lucille F. Stickel Box Turtle Research Award


In honor of the life and work of Lucille F. Stickel, the North American Box Turtle Conservation Committee has established the Lucille F. Stickel Box Turtle Research Award. This award supports conservation projects and research on North American Box Turtles in the following categories: natural history, ecology, conservation, genetics, reproduction, and epidemiology. The Committee awards one or more grants following a competitive proposal review process. Grants are small, and generally do not exceed $1500.

An application form is available here. To make a tax-deductible donation to support this award, please send an email to We would be happy to provide you with convenient options for donations.

The Lucille F. Stickel Box Turtle Research Award Recipients



Box Turtle Radio Telemetry Research and Education Project

Kathryn Royall & Heidi Dull, Environmental Education, Haw River State Park, Browns Summit, NC (,


Determining Population Estimates of Eastern Box Turtles at two City of Raleigh Nature Preserves

Sara Steffen, Durant Nature Preserve, Raleigh, NC & Horseshoe Farm Nature Preserve, Wake Forest, NC (


Identification of Ranavirus infection-associated genes in a zoo population of Eastern Box Turtles (Terrapene c. carolina)

Steven JA Kimble & Noelle Jurcak, Dept. of Biological Sciences, Towson University, Towson, MD (,


Behavior, ecology, and morphology of the Gulf Coast Box Turtle in Mississippi

Evan Grimes, University of Southern Mississipppi (


Radio-Telemetry Tracking of Confiscated and Non-Confiscated Florida Box Turtles (Terrapene carolina bauri) on a Southwest Florida Barrier Island

Chris Lechowicz & Mike Mills, Sanibel-Captiva Conservation Foundation, Sanibel, FL


The Population Status, Demography and Habitat Use of Florida Box Turtles (Terrapene carolina bauri) in a Coastal Ecosystem

Jordan Donini , Katherine Ebaugh & Adrian Rodriguez, Florida SouthWestern State College, Naples, FL (



Ecology and Behavior of the Gulf Coast Box Turtle (Terrapene carolina major) in the Panhandle of Florida (link)

Jessica Meck1,2, Michael T. Jones2, Lisabeth L. Willey1,2 & Jonathan Mays3. 1Antioch University New England, Keene, NH; 2American Turtle Observatory, New Salem, MA; 3Florida Fish and Wildlife Conservation Commission, Gainesville, FL (


Hybridization and Introgression in North American Box Turtles from the Southeastern United (link)

Bradley T. Martin, Marlis R. Douglas, & Michael E. Douglas, University of Arkansas, Fayetteville, AR



The Maryland Zoo Eastern Box Turtle Monitoring and Citizen Science Program (link)

Katharine Mantzouris, Kevin Barrett, & Ellen Bronson. Conservation Department, The Maryland Zoo, Baltimore, MD (,,


Effects of climate variability on the somatic growth of eastern box turtles: Insights from long-term growth chronologies derived from tree-ring analysis (link)

Kiyoshi Sasaki & Ashley Graham, Biology Department, Winthrop University, Rock Hill, SC (



The Risk and Response of Eastern Box Turtles (Terrapene carolina) to Prescribed Fire in the North Carolina Sandhills

Co-Principal Investigators:

   Kristoffer H. Wild, Center of Excellence for Field Biology, Department of Biology, Austin Peay State University Clarksville, Tennessee

   John H. Roe, Department of Biology, University of North Carolina at Pembroke, Pembroke, North Carolina (


Final Report

Community-based Ecological Research and Conservation of the Yucatán box turtle in Yucatán, Campeche, and Quintana Roo

Co-Principal Investigators:

   Michael T. Jones, Ph.D. and Lisabeth L. Willey, Ph.D., Department of Environmental Conservation, University of Massachusetts, Amherst, MA 01003 (,

   Thomas S.B. Akre, Ph.D. and Erika Gonzalez, Smithsonian Mason School of Conservation, Smithsonian Conservation Biology Institute, Front Royal, VA 22630 (

   Dr. Rodrigo Macip-Rios, University of Puebla, Puebla, Mexico, (


Hatchling movements, behavior, and early age class vulnerability of eastern box turtles to land management including prescribed fire

Lead Principal Investigator:

  Dr. Gary Roloff, Associate Professor Michigan State University Department of Fisheries and Wildlife (


Co-Principal Investigator:

  Tracy Swem, Graduate Research Assistant, MSU Fisheries and Wildlife (


Nest success and neonate ecology of the eastern box turtle Terrapene carolina carolina in the Northern Lower Peninsula, Michigan

Principal Investigator:

   Jennifer A. Moore, Assistant Professor of Biology, Grand Valley State University, Allendale, Michigan (



   Patrick B. Laarman, Biology Graduate Research Assistant, Grand Valley State University,

   Paul W. Keenlance, Assistant Professor of Biology, Grand Valley State University, (


Agency Collaborator:

   Patricia O’Connell, Wildlife Biologist, Cadillac-Manistee Ranger District, Huron-Manistee National Forests,
USDA Forest Service (


Nesting site search and home range of gravid and non-females of Terrapene coahuila

Co-Principal Investigators:

   Biol. Miriam Alejandra Cueto Mares and Biol. Sara Isabel Valenzuela Ceballos, Universidad Juárez del Estado de Durango, Facultad de Ciencias biológicas (





Survivorship and movements of head-started hatchling box turtles (Terrapene carolina) salvaged from displaced and injured females

Lead Principal Investigator:

   Kimberly M. Andrews, Ph.D., Research Coordinator, Georgia Sea Turtle Center (GSTC), Jekyll Island Authority, Jekyll Island, GA (


Co-Principal Investigators:

   Terry M. Norton, DVM, Director of Wildlife Health and Director, GSTC  (

   Joseph Colbert, Research Technician, GSTC, (


Final Report, Poster

Molecular characterization of eastern box turtle (Terrapene carolina carolina) breeding biology

Lead Principal Investigator:

   Rod N. Williams, Ph.D., Assistant Professor of Wildlife Science, Department of Forestry and Natural Resources, Purdue University, West Lafayette, IN (


Co-Principal Investigators:
   Steven J.A. Kimble, Ph.D. candidate, Department of Forestry and Natural Resources, Purdue University  (
   Russell L. Burke, Ph.D., Associate Professor,  Department of Biology, Hofstra University, Uniondale, NY (


Turtles and Telemetry: teaching science to high school students through first hand experience

Principal Investigators:
   Kurt Whitford, M.S., Science teacher, Glen Este High School, Cincinnati, OH (
   Jon Souders, M.S., Science teacher, Glen Este High School, (




Dr. John S. Placyk, Jr., Assistant Professor of Biology, Department of Biology, University of Texas at Tyler, 3900 University Blvd., Tyler, Texas 75799

Title: Conservation Genetics of the North American Box Turtles (Terrapene sp.)

Progress Report (February 2011)
Funding provided by the Lucille F. Stickel Box Turtle Research Award has supported my work on the phylogenetics of the Terrapene genus. It has provided for funding to acquire tissue samples, extract DNA from those tissue samples, amplify and sequence genes, and analyze the sequence data. To date, via a vast network of volunteers, we have collected ~1500 tissue samples from every currently recognized subspecies/species of Terrapene with the exception of T. nelsoni; more samples continue to be received on a regular basis. All of the samples in our possession have been inventoried and most have had their DNA extracted. In addition, we are very close to having mtDNA sequence data for every species and subspecies, and most of the states within the North American box turtle distribution are already represented in the mtDNA dataset.

The current mtDNA dataset based on the cytochrome b gene indicates a variety of relationships. To begin, it is clear that there are two main groups (an eastern and a western). However, relationships within those groups appear a bit more muddied. To begin, the data indicate that the three-toed box turtle (T. carolina triunguis) is more distantly related to the eastern box turtle (T. c. carolina) than previously believed (Figure 1). In fact, they may be distinct enough that T. c. triunguis may need to be elevated to a separate species. In addition, our mtDNA dataset indicates that the ornate box turtle (T. o. ornata) are less distantly related to the desert box turtles (T. o. luteola) than previously believed (Figure 1) and that the two may be safely folded into one single species rather than two separate subspecies. Additional evolutionary relationships that appear to be muddled include the placement of the gulf coast, Mexican, and Florida box turtles, as their distribution in the dataset is not straight forward.

Since the current phylogeny is based solely on mtDNA data, we are in the process of acquiring additional data from nuclear sequences to confirm the relationships revealed so far. We expect to have a full data set on one nuclear gene completed within the next month and will sequence at least one more prior to preparing our data for publication. At that time, we will be sure to acknowledge funding from the Lucille F. Stickel Box Turtle Research Award and will forward a copy of the accepted manuscript to the board. Thank you so much for your support!!!


Jeanne M. Baker, graduate student at the University of Illinois, The Illinois Natural History Survey, 1816 South Oak Street, Champaign, IL 61820

Title: The Nesting Ecology of the Eastern Box Turtle, Terrapene carolina at Kickapoo State Park/ Middlefork State Recreation Area


The most researched aspects of Terrapene reproductive ecology cover mating behaviors and egg production (Dodd 2001). Nesting ecology and nest success, however, are largely understudied and past studies have focused mostly on the Ornate Box Turtle, Terrapene ornata (Converse et al . 2002). It is likely that the lack of box turtle nesting data is a result of unpredictable nesting behaviors.

The nesting of T. carolina can occur between early May and the middle of July (Legler 1960; Dodd 2001). The time of day at which nesting occurs varies depending on temperature and other weather conditions. Terrapene carolina will generally nest just after, before, or during a rainfall (Congello 1978; Dodd 2001). The duration of nesting activities is highly variable; sometimes lasting up to five hours, and likely depends on environmental conditions (Ewing 1935; Stickel 1950; Legler 1960; Dodd 2001).

Nest site selection influences the development and survival of young turtles (Dodd 2001; Kolbe and Janzen 2001). Substrate type, temperature, and moisture dictate hatching success. Prior to oviposition, female T. carolina move sporadically around the general nesting site (Stickel 1989). Between one and seven eggs per clutch are deposited, depending on the population's geographic location (Dodd 2001). For example, northern populations deposit four or more eggs and southern populations deposit one to three eggs (Ernst et al . 1994; Dodd 2001). Geographic location and temperature will also influence the emergence time of young turtles. The field incubation period is approximately 65 days, however warmer temperatures may increase development rate, causing eggs to hatch sooner (Dodd 2001).


The study site was located in East Central Illinois, along the Collison Branch of the Middle Fork Vermilion River . Beginning in the spring of 2009, box turtles were located and captured through visual encounter surveys. Each individual was uniquely marked by notching marginal scutes (Cagle 1939). Individuals were weighed on an electronic balance and aged by counting the annuli on their left pectoral scute (Sexton 1959). Carapace length and width, shell height, left pectoral scute length, anterior plastron length and width, and posterior plastron length and width were measured. Adult females were x-rayed to determine clutch sizes (Gibbons and Greene 1979). A small radio transmitter was glued to the carapace of each individual. Individuals were tracked daily and the following environmental data was measured: relative humidity, wind speed, air temperature, substrate temperature, and percent canopy cover. GPS coordinates were also taken.


In the spring of 2009, nine female Terrapene carolina were found to be gravid. The mean clutch size was 5.6 eggs (Table 1). Though females were monitored daily, none were observed nesting. The lack of nesting data from my recent findings is disappointing; however, a summary of my results are as follows:

Search Effort

The search effort consisted of 205 hours between the dates April 23 and June 10. The 47 individuals found between these dates (21 females, 23 males, 3 immature), resulted in an overall catch per unit effort of 0.23 turtles/search hour. The catch per unit effort for gravid females was not determined; only 12 adult females were x-rayed. Searches for individuals continued until October 28 and a total of 95 individuals were found at the study site (36 females, 54 males, 5 immature). The resulting density estimate for 2009 was approximately 0.89 turtles/ha.

Clutch Sizes

In 2009, 12 female Terrapene carolina found before June 9th were x-rayed. Only nine of these individuals were gravid. Three individuals had a clutch size of 7 eggs (1L 2R, 3L 3R, 9L 8R), two had a clutch size of 6 eggs (10L 11L, 3L 11L), one individual had a clutch size of 5 eggs (1L 1R), and three individuals had a clutch size of 4 eggs (1L 12L, 1L 11L, 9L 9R).


The x-rayed individuals were never observed nesting. A single, unmarked female; however, was observed covering a nest at 19:24 on June 13. The female nested on a gravel road near the entrance to the field site. The nest was approximately 2 meters from a wooded edge. The air temperature was 24.5 degrees Celsius and the relative humidity was 85%. The substrate was a mix of dirt and gravel. At the time of nesting, the substrate was 22.2 degrees Celsius (Table 2). The nest was never depredated and hatching was not observed. Nesting movements were not analyzed because the nesting observation was incidental and the female was not fit with a radio transmitter.


My disappointing results obviously preclude in depth analysis of the original objectives of this project. Several factors contributed to these substandard results:

Although the totaling number of female Terrapene carolina was 36 individuals, the number of mature females found before the start of the nesting season indicated that obtaining an adequate sample size of gravid females may not be possible. Females found 1-2 days before the scheduled x-ray session (12 individuals) were photographed. After nine gravid females were returned to their habitat, they immediately moved long distances within the site. It took several days to re-locate these individuals and it is likely that the nesting events took place before the individuals were found.

Terrapene carolina utilized a much larger portion of the study site than expected. The active area of the site was approximately 107 hectares. The habitat was very diverse and difficult to navigate. Private land , ravines, and old fence rows divided the site into four main habitats; grasslands, wet meadows, upland forests, and floodplain forests. It was concluded that navigating through the site during evening hours would be too dangerous.

Field help was a limiting factor for this project. The daily locating of radioed individuals took much longer than expected. Originally, I had planned on monitoring gravid females twice daily (once in the morning and once in the evening). Tracking individuals once during the day would sometimes take up to 10 hours even with field help.

Although the study was not carried out according to the original project outline, I was able to adjust the objectives and managed to retrieve some interesting and valuable data. Remaining funds were used to monitor both male and female Terrapene carolina to study the home range, movement, and habitat use of these individuals. Data from 2009 was presented in the form of a written thesis and submitted to the graduate school at the University of Illinois.

This project was the start of a long-term study in which home range, density estimates, habitat use, and hibernation will be considered. Upon the completion of my graduate work, I continued to locate radioed individuals until hibernation. Each hibernation site was set with a data logger which prepared for the continuation of this study.


Cagle, F.R. 1939. A system for marking turtles for future identification. Copeia 1939:170-173.

Congello, K. 1978. Nesting and egg laying behavior in Terrapene carolina . Proceedings of the Pennsylvania Academy of Science 52:51-56.

Converse, S.J., Iverson J.B., and J.A. Savidge. 2002. Activity, reproduction and overwintering behavior of ornate box turtles ( Terrapene Ornata Ornata ) in the Nebraska sandhills. American Midland Naturalist. 148 (2):416-422.

Dodd, C.K., Jr. 2001. North American box turtles: A natural history. Norman : University of Oklahoma Press, 1-231

Ernst, C.H., Lovich, J.E., and R.W. Barbour. 1994 Turtles of the United States and Canada . Washington and London : Smithsonian Institute Press, 1-578.

Ewing , H.E. 1935. Further notes on the reproduction of the eastern box turtle, Terrapene carolina . Copeia 1935: 102.

Gibbons, J. W. and J.L. Greene. 1979. X-ray photography: A technique to determine reproductive patterns of freshwater turtles. Herpetologica. 35 (1):86-89.

Kolbe, J.J., and F.J. Janzen. 2001. The influence of propagule size and maternal nest-site selection on survival and behaviour of neonate turtles. Functional Ecology. 15 (6):772-781.

Legler, J.M. 1960. Natural history of the ornate box turtle, Terrapene ornata ornata Agassiz . University of Kansas Publications Museum of Natural History. 11: 527-669.

Sexton, O.J. 1959. Spatial and temporal movements of a population of the painted turtle, Chrysemys picta marginata ( Agassiz ). Ecological Monographs. 29:113-140.

Stickel, L.F. 1950. Populations and home range relationships of the box turtle, Terrapene c. carolina (Linnaeus). Ecological Monographs. 20:351-378.

Stickel, L.F. 1989. Home range behavior among box turtles ( Terrapene c. carolina ) of a bottomland forest in Maryland . Journal of Herpetology. 23:40-41.