Timothy S. Mitchell
Iowa State University
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Featured researches published by Timothy S. Mitchell.
Ecology | 2013
Timothy S. Mitchell; Daniel A. Warner; Fredric J. Janzen
Identifying the relative contributions of genetic, maternal, and environmental factors to phenotypic variation is critical for evaluating the evolutionary potential of fitness-related traits. We employed a novel two-step cross-fostering experiment to quantify the relative contributions of clutch (i.e., maternal identity) and maternally chosen nest sites to phenotypic variation during three early life stages (incubation, hibernation, dispersal) of the painted turtle (Chrysemys picta). By translocating eggs between nests in the field, we demonstrated that both clutch and nest site contribute to phenotypic variation at hatching. Because hatchling C. picta hibernate inside nests, we performed a second cross-foster to decouple the effects of the incubation nest with that of the hibernation nest. Incubation nest explained little variation in phenotypes at spring emergence, but winter nest site was important. We found no evidence that mothers select nest sites specific to reaction norms of their own offspring, suggesting that females may select nest sites with microhabitats that broadly meet similar requirements across the population. After hibernation, we released hatchlings to assess performance and phenotypic selection during dispersal. Hibernation nest site influenced physiological performance during dispersal, and we detected nonlinear selection on hatchling carapace length. Our experiment demonstrates that nest-site choice has substantial effects on phenotypic variation and fitness across multiple early life stages.
Proceedings of the Royal Society of London B: Biological Sciences | 2013
Timothy S. Mitchell; Jessica A. Maciel; Fredric J. Janzen
Evolutionary theory predicts that dioecious species should produce a balanced primary sex ratio maintained by frequency-dependent selection. Organisms with environmental sex determination, however, are vulnerable to maladaptive sex ratios, because environmental conditions vary spatio-temporally. For reptiles with temperature-dependent sex determination, nest-site choice is a behavioural maternal effect that could respond to sex-ratio selection, as mothers could adjust offspring sex ratios by choosing nest sites that will have particular thermal properties. This theoretical prediction has generated decades of empirical research, yet convincing evidence that sex-ratio selection is influencing nesting behaviours remains absent. Here, we provide the first experimental evidence from nature that sex-ratio selection, rather than only viability selection, is probably an important component of nest-site choice in a reptile with temperature-dependent sex determination. We compare painted turtle (Chrysemys picta) neonates from maternally selected nest sites with those from randomly selected nest sites, observing no substantive difference in hatching success or survival, but finding a profound difference in offspring sex ratio in the direction expected based on historical records. Additionally, we leverage long-term data to reconstruct our sex ratio results had the experiment been repeated in multiple years. As predicted by theory, our results suggest that sex-ratio selection has shaped nesting behaviour in ways likely to enhance maternal fitness.
Functional Ecology | 2015
Timothy S. Mitchell; Jessica A. Maciel; Fredric J. Janzen
Summary Offspring phenotypic variation can be substantially influenced by non-genetic factors such as maternal effects, which ultimately can influence organismal fitness. For oviparous organisms that lack parental care, oviposition-site choice and egg size are maternal effects that can greatly affect offspring traits. Yet, few studies examine the consequences of these traits in the wild. We manipulated the contents of natural painted turtle nests such that offspring spent two life stages (incubation and hibernation) in either maternally selected nest sites or randomly selected nest sites and quantified treatment differences in environmental parameters and offspring phenotypes. Additionally, we tracked the fates of individual eggs and hatchlings, which allowed us to quantify the strength and form of selection acting on egg size during incubation and, for the first time, body size during hibernation. Maternally selected nest sites were warmer and produced offspring that were longer and hatched earlier than their siblings emerging from cooler, randomly selected nests. Treatments did not affect any measured traits during hibernation. We detected no selection on egg size during the incubation stage, but significant linear selection favouring larger hatchlings during hibernation. Our results suggest that nest-site choice allows mothers to partially control the environment of their incubating eggs, but is less effective at controlling hatchling environments during hibernation. Additionally, we provide novel support for the ‘bigger-is-better’ hypothesis in turtles by showing a positive relationship between size and survival during the hibernation stage.
The American Naturalist | 2016
Rory S. Telemeco; Eric J. Gangloff; Gerardo A. Cordero; Timothy S. Mitchell; Brooke L. Bodensteiner; Kaitlyn G. Holden; Sarah M. Mitchell; Rebecca L. Polich; Fredric J. Janzen
Historically, egg-bound reptile embryos were thought to passively thermoconform to the nest environment. However, recent observations of thermal taxis by embryos of multiple reptile species have led to the widely discussed hypothesis that embryos behaviorally thermoregulate. Because temperature affects development, such thermoregulation could allow embryos to control their fate far more than historically assumed. We assessed the opportunity for embryos to behaviorally thermoregulate in nature by examining thermal gradients within natural nests and eggs of the common snapping turtle (Chelydra serpentina; which displays embryonic thermal taxis) and by simulating thermal gradients within nests across a range of nest depths, egg sizes, and soil types. We observed little spatial thermal variation within nests, and thermal gradients were poorly transferred to eggs. Furthermore, thermal gradients sufficiently large and constant for behavioral thermoregulation were not predicted to occur in our simulations. Gradients of biologically relevant magnitude have limited global occurrence and reverse direction twice daily when they do exist, which is substantially faster than embryos can shift position within the egg. Our results imply that reptile embryos will rarely, if ever, have the opportunity to behaviorally thermoregulate by moving within the egg. We suggest that embryonic thermal taxis instead represents a play behavior, which may be adaptive or selectively neutral, and results from the mechanisms for behavioral thermoregulation in free-living stages coming online prior to hatching.
Functional Ecology | 2015
Brooke L. Bodensteiner; Timothy S. Mitchell; Jeramie T. Strickland; Fredric J. Janzen
Summary 1. Phenotypic variation is strongly impacted by environmental conditions experienced during development. Substantial laboratory research has shown that reptiles with flexible-shelled eggs are particularly sensitive to hydric conditions, yet research on nests in the wild is sparse. 2. In this 2-year field experiment, we explore the influence of hydric conditions during incubation on phenotypic traits of hatchling painted turtles (Chrysemys picta). Using a split-clutch design, we created two artificial nests adjacent to each maternally selected nest site. Half the eggs incubated in a nest that received regular supplemental watering, while the control nest was exposed to natural precipitation only. 3. Our results suggest that the influence of the hydric environment on developing reptilian embryos is context dependent. Supplemental water applied to nests in a drier than normal season elicited the expected biotic responses, based on laboratory experiments. However, when the soil surrounding C. picta eggs was already highly moist, the additional water from supplemental application effectively stunted embryonic development. 4. Our experiment confirms that hydric conditions of the soil during incubation in the wild can substantially influence phenotypic variation of reptiles with flexible-shelled eggs. Additionally, our experiment highlights the importance of complex interactions in the field that are often unexplored in laboratory experiments, reiterating the importance of validating laboratory work with field experiments.
Oecologia | 2013
Daniel A. Warner; Timothy S. Mitchell
Orientation and dispersal to suitable habitat affects fitness in many animals, but the factors that govern these behaviors are poorly understood. In many turtle species, hatchlings must orient and disperse to suitable aquatic habitat immediately after emergence from subterranean nests. Thus, the location of nest sites relative to aquatic habitats ideally should be associated with the direction of hatchling dispersal. At our study site, painted turtles (Chrysemys picta) nest to the west (on an island) and east (on the mainland) of a wetland, which determines the direction that hatchlings must travel to reach suitable aquatic habitat. To determine if hatchling orientation is intrinsically influenced by the location where their mothers nest, we employed a two-part cross-fostering experiment in the field, whereby half the eggs laid in mainland nests were swapped with half the eggs laid in island nests. Moreover, because C. picta hatchlings overwinter inside their nests, we performed a second cross-fostering experiment to fully decouple the effects of (1) the maternally chosen nest location, (2) the embryonic developmental location, and (3) the overwinter location. We released hatchlings into a circular arena in the field and found that turtles generally dispersed in a westerly direction, regardless of the maternally chosen nest location and independent of the locations of embryonic development and overwintering. Although this westerly direction was towards suitable aquatic habitat, we could not distinguish whether naïve hatchling turtles (i) use environmental cues/stimuli to orient their movement, or (ii) have an intrinsic bias to orient west in the absence of stimuli. Nevertheless, these findings suggest that the orientation behavior of naïve hatchling turtles during terrestrial dispersal is not dependent upon the location of maternally-chosen nest sites.
Journal of Experimental Zoology | 2017
Austin C. Hulbert; Timothy S. Mitchell; Joshua M. Hall; Cassia M. Guiffre; Danielle C. Douglas; Daniel A. Warner
Many studies of phenotypic plasticity alter environmental conditions during embryonic development, yet only measure phenotypes at the neonatal stage (after embryonic development). However, measuring aspects of embryo physiology enhances our understanding of how environmental factors immediately affect embryos, which aids our understanding of developmental plasticity. While current research on reptile developmental plasticity has demonstrated that fluctuating incubation temperatures affect development differently than constant temperatures, most research on embryo physiology is still performed with constant temperature experiments. In this study, we noninvasively measured embryonic heart rates of the brown anole (Anolis sagrei), across ecologically relevant fluctuating temperatures. We incubated eggs under temperatures measured from potential nests in the field and examined how heart rates change through a diel cycle and throughout embryonic development. We also evaluated how experimental design (e.g., repeated vs. single measures designs, constant vs. fluctuating temperatures) and different protocols (e.g., removing eggs from incubators) might influence heart rate. We found that heart rates were correlated with daily temperature and increased through development. Our findings suggest that experimenters have reasonable flexibility in choosing an experimental design to address their questions; however, some aspects of design and protocol can potentially influence estimations of heart rates. Overall, we present the first ecologically relevant measures of anole embryonic heart rates and provide recommendations for experimental designs for future experiments.
Journal of Experimental Zoology | 2018
Timothy S. Mitchell; Fredric J. Janzen; Daniel A. Warner
Studies of reptiles have contributed greatly to understanding the impacts of developmental environments on offspring phenotypes. A major challenge for these studies, however, is quantifying the effects of embryonic environments on adult phenotypes and reproductive success. Such measurements may be necessary to gain full insight into the evolution of plasticity, as well as the long-term consequences of plasticity under environmental change. Unfortunately, most studies of reptile developmental plasticity only measure phenotypic traits of offspring at hatching, and rarely evaluate effects on subsequent adult phenotypes. This lack of information highlights a major gap in this active field. In this review, we first discuss conceptual issues regarding the ecology and evolution of plasticity to provide justification for long-term studies necessary to measure adult phenotypes. Second, we review case studies of reptiles that assessed the effects of developmental environments on adult phenotypes and/or reproduction, and we highlight the valuable insights that they provide. Importantly, we illustrate that terminating studies during early-life stages can lead to incomplete or even misleading interpretations. Third, we discuss the pros and cons of different experimental approaches for quantifying long-term effects of developmental environments. Overall, devoted long-term studies on taxa with diverse ecologies and life histories will provide major advances in the field of developmental plasticity.
Journal of Experimental Zoology | 2017
Daniel A. Warner; Timothy S. Mitchell; Brooke L. Bodensteiner; Fredric J. Janzen
Exogenous application of steroids and related substances to eggs affects offspring sex ratios in species with temperature-dependent sex determination (TSD). Laboratory studies demonstrate that this effect is most pronounced near the constant temperature that produces 1:1 sex ratios (i.e., pivotal temperature). However, the impact of such chemicals on sex determination under natural nest temperatures (which fluctuate daily) is unknown, but could provide insight into the relative contributions of these two factors under natural conditions. We applied estradiol (E2) and an aromatase inhibitor (fadrozole) to eggs of the painted turtle (Chrysemys picta), a species with TSD, and allowed eggs to incubate under natural conditions during two field seasons (in 2012 and 2013). Exogenous E2, fadrozole, and nest temperature contributed to variation in offspring sex ratio, but the relative contributions of these factors differed between years. In 2012, a much hotter than average season, sex ratios were heavily female biased regardless of nest temperature and chemical treatment. However, in 2013, a milder season, both nest temperature and chemical treatment were important. Moreover, a significant interaction between nest temperature and treatment demonstrated that exogenous estradiol induces female development regardless of nest temperature, but aromatase inhibition widens the range of temperatures that produces both sexes.
American Midland Naturalist | 2016
Kameron C. Voves; Timothy S. Mitchell; Fredric J. Janzen
Abstract Egg and nest crypsis is a strategy to reduce detection by predators, thereby minimizing offspring mortality. While this strategy has been well studied in birds, it has received little attention in other taxa. Turtles are plausibly able to camouflage their subterranean nests by reducing the level of soil surface disturbance. To test the hypothesis that more cryptic nests experience lower predation, we first quantified the camouflage of natural Painted Turtle (Chrysemys picta) nests from the human perspective and demonstrated that substantial variation in nest camouflage exists. We subsequently tracked the predation fates of these nests and related nest camouflage to nest survival. Although the results trend in the expected direction, logistic regression did not yield a significant association between camouflage rating and nest survival. These results suggest turtle nests are highly detectable by predators and mothers have little ability to influence predation risk to their nests via camouflage. This work opens a previously little-explored area in nest crypsis in which further study could aid in discerning a functional connection between turtle nest camouflage and survival.