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Todd Lab Herpetolgy and Wildlife Conservation

Todd Lab publications

The following is a list of publications by members of our lab (members indicated in bold). Many of the publications are available as PDF downloads by clicking the title of the article or are available at the links under open access licenses.


Glass MS, Todd BD, Buhlmann KA, Rushing CS, Barnas G, Tuberville TD. 2024. Survival and behavior of Mojave desert tortoises head-started with and without outdoor rearing. Journal of Wildlife Management 88:e22562.

Nguyen AM, Halstead BJ, Todd BD. 2024. Effect of translocation on home range and movements of giant gartersnakes. Global Ecology and Conservation 49:e02789.

Richter CJ, Todd BD, Buhlmann KA, Candal CM, McGovern, PA, Kohl MT, Tuberville TD. 2024. Effects of head-starting on multi-year space use and survival of an at-risk tortoise. Global Ecology and Conservation 49:e02774.


Nowakowski AJ, Watling JI, Murray A, Deichmann JL, Akre TA, Muñoz-Brenes CL, Todd BD, McRae L, Freeman R, Frishkoff LO. 2023. Protected areas slow declines unevenly across the vertebrate tree of life. Nature 622:101–106. (Email me for copy)

Forrest MJ, Halstead BJ, Grear DA, Kleeman PM, Todd BD, Miano OJ Urquhart KD. 2023. Keeping the heat on: Weighted surveillance for chytrid fungus in Dixie Valley toads. Journal of Wildlife Diseases 59(4):557–568.

Nguyen AM, Todd BD, Halstead BJ. 2023. Survival and establishment of captive‐reared and translocated giant gartersnakes after release. Journal of Wildlife Management 87(3):e22374.


Todd BD, Jenkinson TS, Escalona M, Beraut E, Nguyen O, Sahasrabudhe R, Scott PA, Toffelmier E, Wang IJ, Shaffer HB. 2022. Reference genome of the northwestern pond turtle, Actinemys marmorata. Journal of Heredity 113(6):624–631.

Greenwood JF, Lara Granados G, Secor SM, Todd BD, Showalter I, Hedrick BP, Brennan PLR. 2022. Divergent genital morphologies and female–male covariation in water snakes. Integrative and Comparative Biology 62(3):569–580.

Nafus MG, Daly JA, Tuberville TD, Klimley AP, Buhlmann KA, Todd BD. 2022. Habitat use by desert tortoises suggests tradeoffs between resource use and risk avoidance. PLoS ONE 17(8):e0263743.


Brown C, Nowakowski AJ, Keung NC, Lawler SP, Todd BD. 2021. Untangling multi-scale habitat relationships of an endangered frog in streams to inform reintroduction programs. Ecosphere 12(10):e03799.

McGovern PA, Peaden JM, Buhlmann KA, Todd BD, Tuberville TD. 2021. Comparing post-release cover and burrow use by differentially head-started Mojave desert tortoises in southern California. Conservation Evidence 18:37–43.

Keung NC, Lawler SP, Yarnell SM, Todd BD, Brown C. 2021. Movement ecology study of stream-dwelling Sierra Nevada Yellow-legged Frogs (Rana sierrae) to inform reintroductions. Herpetological Conservation and Biology 16(1):72–85.

Todd BD, Nowakowski AJ. 2021. Ectothermy and the macroecology of home range scaling in snakes. Global Ecology and Biogeography 30(1):262–276. --[Data available here: Data Dryad Repository ]


McGovern PA, Buhlmann KA, Todd BD, Moore CT, Peaden JM, Hepinstall-Cymerman J, Daly JA, Tuberville TD. 2020. The effect of size on postrelease survival of head-started Mojave Desert Tortoises. Journal of Fish and Wildlife Management 11(2):494–506.

McGovern PA, Buhlmann KA, Todd BD, Moore CT, Peaden JM, Hepinstall-Cymerman J, Daly JA, Tuberville TD. 2020. Comparing husbandry techniques for optimal head-starting of the Mojave Desert Tortoise (Gopherus agassizii). Herpetological Conservation and Biology 15(3):626–641.

Brown C, Keung NC, Dillingham CP, Mussulman S, Bushell J, Sollmann R, Todd BD, Lawler SP. 2020. Using demography to evaluate reintroductions for conservation of the endangered frog, Rana sierrae, in streams. Herpetologica 76(4):383-395.

Rose JP, Todd BD. 2020. Targeting eradication of non-native species using integral projection models. Animal Conservation 23(6):713-724.

Agha M, Yackulic CB, Riley MK, Peterson B, Todd BD. 2020. Brackish tidal marsh management and the ecology of a declining freshwater turtle. Environmental Management 66:644–653.

Agha M, Lovich JE, Ennen JR, Todd BD. 2020. Wind, sun, and wildlife: do wind and solar energy development “short-circuit” conservation in the western United States? Environmental Research Letters 15(7):075004.

Nowakowski AJ, Peaden JM, Tuberville TD, Buhlmann KA, Todd BD. 2020. Thermal performance curves based on field movements reveal context-dependence of thermal traits in a desert ectotherm. Landscape Ecology 35:893–906.


Brown C, Wilkinson LR, Wilkinson KK, Tunstall T, Foote R, Todd BD, Vredenburg VT. 2019. Demography, habitat, and movements of the Sierra Nevada Yellow-legged Frog (Rana sierrae) in streams. Copeia 107(4):661–675.

Yarnell SM, Peek RA, Keung N, Todd BD, Lawler S, Brown C. 2019. A lentic breeder in lotic waters: Sierra Nevada Yellow-Legged Frog (Rana sierrae) habitat suitability in northern Sierra Nevada streams. Copeia 107(4):676–693.

Daly JA, Buhlmann KA, Todd BD, Moore CT, Peaden JM, Tuberville TD. 2019. Survival and movements of head-started Mojave Desert Tortoises. Journal of Wildlife Management 83(8):1700–1710.

Rivera-Ordonez JM, Nowakowski AJ, Manansala A, Thompson ME, Todd BD. 2019. Thermal niche variation among individuals of the poison frog, Oophaga pumilio, in forest and converted habitats. Biotropica 51:747–756.

Agha M, Yanagitsuru YR, Fangue NA, Nowakowski AJ, Kojima LV, Cech JJ, Riley MK, Freeman J, Cocherell DE, Todd BD. 2019. Physiological consequences of rising water salinity for a declining freshwater turtle. Conservation Physiology 7(1):coz054.

Rose JP, Wademan C, Weir S, Wood JS, Todd BD. 2019. Traditional trapping methods outperform eDNA sampling for introduced semi-aquatic snakes. PLoS ONE 14(7):e0219244.

Martinez A, Palumbo S, Todd BD. 2019. Bio-inspiration for anisotropic load transfer at soil-structure interfaces. Journal of Geotechnical and Geoenvironmental Engineering 145(10):04019074.

Tuberville TD, Buhlmann KA, Sollmann R, Nafus MG, Peaden JM, Daly JA, Todd BD. 2019. Effects of short-term, outdoor head-starting on growth and survival in the Mojave Desert Tortoise (Gopherus agassizii). Herpetological Conservation and Biology 14(1):171–184.


Daly JA, Buhlmann KA, Todd BD, Moore CT, Peaden JM, Tuberville TD. 2018. Comparing growth and body condition of indoor-reared, outdoor-reared, and direct-release juvenile Mojave desert tortoises. Herpetological Conservation and Biology 13(3)622–633.

Nowakowski AJ, Frishkoff LA, Agha M, Todd BD, Scheffers BR. 2018. Changing thermal landscapes: merging climate science and landscape ecology through thermal biology. Current Landscape Ecology Reports 3(4):57–72.

Nowakowski AJ, Frishkoff LA, Thompson ME, Smith TM, Todd BD. 2018. Phylogenetic homogenization of amphibian assemblages in human-altered habitats across the globe. Proceedings of the National Academy of Sciences, USA 115:E3454–E3462.

Agha M, Ennen JR, Bower D, Nowakowski AJ, Sweat S, Todd BD. 2018. Salinity tolerances and use of saline environments by freshwater turtles: implications of sea level rise. Biological Reviews 93:1634–1648.

Agha M, Todd BD, Augustine B, Lhotka JM, Fleckensten LJ, Lewis M, Patterson C, Stringer J, Price SJ. 2018. Effects of gap-based silviculture on thermal biology of a terrestrial reptile. Wildlife Research 45:72–81.

Agha M, Riley MK, Sanford E, Carlton JT, Newman WA, Todd BD. 2018. A review of epizoic barnacles reported from freshwater turtles with a new record from California. Herpetological Review 49(1):25–28.

Nowakowski AJ, Watling JI, Brusch G, Catenazzi A, Whitfield SM, Kurz D, Suarez-Mayorga A, Aponte-Gutiérrez A, Donnely MA, Todd BD. 2018. Thermal biology mediates responses of amphibians and reptiles to habitat modification. Ecology Letters 21:345–355.

Eskew EA, Shock BC, LaDouceur EEB, Keel K, Miller MR, Foley JE, Todd BD. 2018. Gene expression differs in susceptible and resistant amphibians exposed to Batrachochytrium dendrobatidis. Royal Society Open Science 5:170910.

Agha M, Ennen JR, Nowakowski AJ, Lovich J, Sweat S, Todd BD. 2018. Macroecological patterns of sexual size dimorphism in turtles of the world. Journal of Evolutionary Biology 31:336–345. [Editor's Choice and cover selection]


Eskew EA, Todd BD. 2017. Too cold, too wet, too bright, or just right? Environmental predictors of snake movement and activity. Copeia 2017:584–591.

Peaden JM, Nowakowski AJ, Buhlmann KA, Tuberville TD, Todd BD. 2017. Effects of roads and roadside fencing on movements, space use, and carapace temperatures of a threatened tortoise. Biological Conservation 214:13–22.

Rose JP, Todd BD. 2017. Demographic effects of prolonged drought on a nascent introduction of a semi-aquatic snake. Biological Invasions 19:2885–2898.

Agha M, Smith AL, Lovich JE, Delaney D, Ennen JR, Briggs J, Fleckensten LJ, Tennant LA, Puffer SR, Walde A, Arundel TR, Price SJ, Todd BD. 2017. Mammalian mesocarnivore visitation at tortoise burrows in a wind farm. Journal of Wildlife Management 81(6):1117–1124.

Nafus MG, Tuberville TD, Buhlmann KA, Todd BD. 2017. Precipitation quantity and timing affect native plant production and growth of a key herbivore, the desert tortoise, in the Mojave Desert. Climate Change Responses 4:4.

Nowakowski AJ, Thompson ME, Donnelly MA, Todd BD. 2017. Amphibian sensitivity to habitat modification is associated with population trends and species traits. Global Ecology and Biogeography 26:700–712.

Agha M, Price SJ, Nowakowski J, Augustine B, Todd BD. 2017. Mass mortality of eastern box turtles with upper respiratory disease following atypical cold weather. Diseases of Aquatic Organisms 124:91–100.

Nowakowski AJ, Watling JI, Whitfield SM, Todd BD, Kurz DJ, Donnelly MA. 2017. Tropical amphibians in shifting thermal landscapes under land use and climate change. Conservation Biology 31(1):96–105.

Todd BD, Nowakowski AJ, Rose JP, Price SJ. 2017. Species traits explaining sensitivity of snakes to human land use estimated from citizen science data. Biological Conservation 206:31–36.


Todd BD, Rose JP, Price SJ, Dorcas ME. 2016. Using citizen science data to identify the sensitivity of species to human land use. Conservation Biology 30(6):1266–1276.

Nowakowski AJ, Whitfield SM, Eskew EA, Thompson ME, Rose JP, Kerby JL, Donnelly MA, Todd BD. 2016. Infection risk decreases with increasing mismatch in host and pathogen environmental tolerances. Ecology Letters 19:1051–1064.

Reed RN, Todd BD, Miano OJ, Canfield MJ, Fisher RN, McMartin L. 2016. Ecology and control of an introduced population of southern watersnakes (Nerodia fasciata) in southern California. Herpetologica 72(2):130–136.

Todd BD, Halstead BJ, Chiquoine LP, Peaden JM, Tuberville TD, Buhlmann KA, Nafus MG. 2016. Habitat selection by juvenile Mojave Desert tortoises. Journal of Wildlife Management 80(4):720–728.

Meyer E, Eskew EA, Chibwe L, Schrlau J, Simonich SLM, Todd BD. 2016. Organic contaminants in western pond turtles in remote habitat in California. Chemosphere 154:326–334.


Peaden JM, Tuberville TD, Buhlmann KA, Nafus MG, Todd BD. 2015. Delimiting road-effect zones for threatened species: implications for mitigation fencing. Wildlife Research 42:650–659.

Rose JP, Miano OJ, Cottle GR, Lovich RE, Palmer RL, Todd BD. 2015. A herpetological inventory of Naval Air Station Fallon, Churchill County, Nevada. Western North American Naturalist 75(4):526–534.

Eskew EA, Worth SJ, Foley JE, Todd BD. 2015. American bullfrogs (Lithobates catesbeianus) resist infection by multiple isolates of Batrachochytrium dendrobatidis, including one implicated in wild mass mortality. EcoHealth 12:513–518.

Nafus MG, Todd BD, Buhlmann KA, Tuberville TD. 2015. Consequences of maternal effects on offspring size, growth, and survival in the desert tortoise. Journal of Zoology 297:108–114.

Zaragoza G, Rose JP, Purcell K, Todd BD. 2015. Terrestrial habitat use by Western Pond Turtles (Actinemys marmorata) in the Sierra foothills. Journal of Herpetology 49(3):437–441.

Nafus MG, Germano JM, Perry JA, Todd BD, Walsh A, Swaisgood RR. 2015. Hiding in plain sight: a study on camouflage and habitat selection in a slow-moving desert herbivore. Behavioral Ecology 26(5):1389–1394.


Tuberville TD, Todd BD, Hermann SM, Michener WK, Guyer C. 2014. Survival, demography, and growth of gopher tortoises (Gopherus polyphemus) from three study sites with different management histories. Journal of Wildlife Management 78(7):1151–1160.

Eskew EA, Todd BD, Hopkins WA. 2014. Extremely low prevalence of Batrachochytrium dendrobatidis infection in Eastern Hellbenders (Cryptobranchus alleganiensis alleganiensis) in Southwest Virginia, USA. Herpetological Review 45(3):425–427.

Rose JP, Todd BD. 2014. Projecting invasion risk of non-native watersnakes (Nerodia fasciataand Nerodia sipedon) in the western United States. PLoS ONE 9(6):e100277. --[Download supplementary North American wetland density GIS layer derived from USGS National Hydrography dataset used in this publication.]

Todd BD, Blomquist SM, Harper EB, Osbourn MS. 2014. Effects of timber harvesting on terrestrial survival of pond-breeding amphibians. Forest Ecology and Management 313:123–131.

Showalter I, Todd BD, Brennan P. 2014. Intraspecific and interspecific variation of female genitalia in two species of watersnakes. Biological Journal of the Linnean Society 111:183–191.


Lambert MR, Yasuda CM, Todd BD. 2013. Ontogenetic shift in habitat use in the western fence lizard (Sceloporus occidentalis) in northern California. Herpetological Review 44(4):579–582.

Rose JP, Miano OJ, Todd BD. 2013. Trapping efficiency, demography, and density of an introduced population of Northern Watersnakes, Nerodia sipedon, in California. Journal of Herpetology 47(3):421–427.

Nafus MG, Tuberville TD, Buhlmann KA, Todd BD. 2013. Relative abundance and demographic structure of Agassiz’s desert tortoise (Gopherus agassizii) along roads of varying size and traffic volume. Biological Conservation 162:100–106.

Eskew EA, Todd BD. 2013. Parallels in amphibian and bat declines from pathogenic fungi. Emerging Infectious Diseases 19(3):379–385.


Willson JD, Hopkins WA, Bergeron CM, Todd BD. 2012. Making leaps in amphibian ecotoxicology: translating individual-level effects of contaminants to population viability. Ecological Applications 22(6): 1791–1802.

Lambert MR, Yasuda CM, Todd BD. 2012. Evaluation of a photographic technique for estimating body size in lizards from a distance. Herpetological Conservation and Biology 7(1):83–88.

Todd BD, Bergeron CM, Hopkins WA. 2012. Use of toe clips as a nonlethal index of mercury accumulation and maternal transfer in amphibians. Ecotoxicology 21:882–887.

Todd BD, Willson JD, Bergeron CM, Hopkins WA. 2012. Do effects of mercury in larval amphibians persist after metamorphosis? Ecotoxicology 21:87–95.


Thornhill DJ, Rotjan RD, Todd BD, Chilcoat GC, Iglesias-Prieto R, Kemp DW, LaJeunesse TC, Reynolds JM, Schmidt GW, Shannon T, Warner ME, Fitt WK. 2011. A connection between colony biomass and death in Caribbean reef-building corals. PLoS ONE 6(12): e29535. doi:10.1371/journal.pone.0029535

Todd BD, Scott DE, Pechmann JHK, Gibbons JW. 2011. Climate change correlates with rapid delays and advancements in reproductive timing in an amphibian community. Proceedings of the Royal Society B 278:2191–2197. Press coverage: (1) Nature (2) Reuters (3) The Ecologist

Wada H, Bergeron CM, McNabb FMA, Todd BD, Hopkins WA. 2011. Dietary mercury has no observable effects on thyroid-mediated processes and fitness-related traits in wood frogs. Environmental Science & Technology 45(18):7915–7922.

Todd BD, Bergeron CM, Hepner MJ, Hopkins WA. 2011. Aquatic and terrestrial stressors in amphibians: a test of the double jeopardy hypothesis using maternally and trophically derived mercury. Environmental Toxicology & Chemistry 30(10):2277–2284.

Todd BD, Bergeron CM, Hepner MJ, Burke JN, Hopkins WA. 2011. Does maternal exposure to an environmental stressor affect offspring response to predators? Oecologia 166(1):283–290.

Bergeron CM, Hopkins WA, Todd BD, Hepner MJ, Unrine JM. 2011. Interactive effects of maternal and dietary mercury exposure have latent and lethal consequences for amphibian larvae. Environmental Science & Technology 45:3781–3787. Press coverage: American Chemical Society

Willson JD, Winne CT, Todd BD. 2011. Ecological and methodological factors affecting detectability and population estimation in elusive species. Journal of Wildlife Management 75(1):36–45.


Burke JN, Bergeron CM, Todd BD, Hopkins WA. 2010. Effects of mercury on behavior and performance of northern two-lined salamanders (Eurycea bislineata). Environmental Pollution 158(12):3546–3551.

Todd BD, Willson JD, Gibbons JW. 2010. The global status of reptiles and causes of their decline. Chapter 3 pp. 47–67. In, Ecotoxicology of Amphibians and Reptiles, 2nd Ed. Sparling, D.W., Linder, G., Bishop, C.A., and Krest, S. (eds) CRC Press, Pensacola, FL, USA.


Semlitsch RD, Todd BD, Blomquist SM, Calhoun AJK, Gibbons JW, Gibbs JP, Graeter GJ, Harper EB, Hocking DJ, Hunter ML, Patrick DA, Rittenhouse TAG, Rothermel, BB. 2009. Effects of timber harvest on amphibian populations: understanding mechanisms from forest experiments. Bioscience 59:853–862. (includes cover photo)

Todd BD, Luhring TM, Rothermel BB, Gibbons JW. 2009. Effects of forest removal on amphibian migrations: implications for habitat and landscape connectivity. Journal of Applied Ecology 46:554–561.


Tuberville TD, Norton TM, Todd BD, Spratt JS. 2008. Long-term apparent survival of translocated gopher tortoises: a comparison of newly released animals and previously established residents. Biological Conservation 141:2690–2697.

Todd BD, Willson JD, Winne CT, Gibbons JW. 2008. Aspects of the ecology of the earth snakes (Virginia valeriae and V. striatula) in the Upper Coastal Plain. Southeastern Naturalist 7(2):349–358.

Todd BD, Andrews KM. 2008. Response of a reptile guild to forest harvesting. Conservation Biology 22(3):753–761.

Todd BD, Willson JD, Winne CT, Semlistch RD, Gibbons JW. 2008. Ecology of the Southeastern Crowned Snake, Tantilla coronata. Copeia 2008(2):388–394.

Todd BD, Rothermel BB, Reed RN, Luhring TM, Schlatter K, Trenkamp L, Gibbons JW. 2008. Habitat alteration increases invasive fire ant abundance to the detriment of amphibians and reptiles. Biological Invasions 10:539–546.


Todd BD, Winne CT, Willson JD, Gibbons JW. 2007. Getting the drift: examining the effects of timing, trap type, and taxon on herpetofaunal drift fence surveys. American Midland Naturalist 158(2):292–305.

Todd BD. 2007. Parasites lost? An overlooked hypothesis for the evolution of alternative reproductive strategies in amphibians. The American Naturalist 170(5):793–799.

Todd BD, Davis AK. 2007. Sexual dichromatism in the marbled salamander, Ambystoma opacum. Canadian Journal of Zoology 85(9):1008–1013.

Winne CT, Willson JD, Todd BD, Andrews KM, Gibbons JW. 2007. Enigmatic decline of a protected population of Eastern Kingsnakes, Lampropeltis getula, in South Carolina. Copeia 2007(3):507–519.

Baughman B, Todd BD. 2007. The role of substrate cues in habitat selection by juvenile Bufo terrestris and Scaphiopus holbrookii. Journal of Herpetology 41(1):154–157.


Todd BD, Rothermel BB. 2006. Assessing quality of clearcut habitats for amphibians: effects on abundances versus vital rates in the southern toad (Bufo terrestris). Biological Conservation 133(2):178–185.

Gibbons JW, Winne CT, Scott DE, Willson JD, Gladaus X, Andrews KM, Todd BD Fedewa LA, Wilkinson L, Tsaliagos RN, Harper SJ, Greene JL, Tuberville TD, Metts BS, Dorcas ME, Nestor JP, Young CA, Akre T, Reed RN, Buhlmann KA, Norman J, Croshaw DA, Hagen C, Rothermel BB. 2006. Remarkable amphibian biomass and abundance in an isolated wetland: implications for wetland conservation. Conservation Biology 20(5):1457–1465.

Todd BD, Winne CT. 2006. Ontogenetic and interspecific variation in timing of movement and responses to climatic factors during migrations by pond-breeding amphibians. Canadian Journal of Zoology 84(5):715–722.

Todd BD, Thornhill DJ, Fitt WK. 2006. Patterns of inorganic phosphate uptake in Cassiopea xamachana: a bioindicator species. Marine Pollution Bulletin 52(5):515–521.