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Creators/Authors contains: "Clark, Rulon"

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  1. Abstract Hybridization between species provides unique opportunities to understand evolutionary processes that are linked to reproductive isolation and, ultimately, speciation. However, the extrinsic factors that limit hybridization are poorly understood for most animal systems. Although the spatial ecology of individuals in natural habitats is fundamental to shaping reproductive success and survival, analyses of the spatial ecology of hybrids and their parental groups are rarely reported. Here, we used radiotelemetry to monitor wild rattlesnakes across an interspecific hybrid zone (Crotalus scutulatus and Crotalus viridis) and measured movement parameters and space use (utilization distributions) of individuals to evaluate the hypothesis that hybridization resulted in transgressive or atypical movement patterns. Unexpectedly, of the spatial metrics we investigated, we found that hybrids were very similar to parental individuals. Nonetheless, hybrids did show increased patchiness of core utilization distributions, but this result is likely to be driven by increased habitat patchiness in the hybrid zone. Overall, we did not find evidence for overt extrinsic barriers to hybridization associated with spatial ecology; thus, we suggest that the close evolutionary history between the two parental species and their ecological and behavioural similarities are likely to increase the probability of hybridization events in this unique region of New Mexico. 
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  2. Abstract Climate change has the potential to disrupt species interactions across global ecosystems. Ectotherm–endotherm interactions may be especially prone to this risk due to the possible mismatch between the species in physiological response and performance. However, few studies have examined how changing temperatures might differentially impact species' niches or available suitable habitat when they have very different modes of thermoregulation. An ideal system for studying this interaction is the predator–prey system. In this study, we used ecological niche modeling to characterize the niche overlap and examine biogeography in past and future climate conditions of prairie rattlesnakes (Crotalus viridis) and Ord's kangaroo rats (Dipodomys ordii), an endotherm–ectotherm pair typifying a predator–prey species interaction. Our models show a high niche overlap between these two species (D = 0.863 andI = 0.979) and further affirm similar paleoecological distributions during the last glacial maximum (LGM) and mid‐Holocene (MH). Under future climate change scenarios, we found that prairie rattlesnakes may experience a reduction in overall suitable habitat (RCP 2.6 = −1.82%, 4.5 = −4.62%, 8.5 = −7.34%), whereas Ord's kangaroo rats may experience an increase (RCP 2.6 = 9.8%, 4.5 = 11.71%, 8.5 = 8.37%). We found a shared trend of stable suitable habitat at northern latitudes but reduced suitability in southern portions of the range, and we propose future monitoring and conservation be focused on those areas. Overall, we demonstrate a biogeographic example of how interacting ectotherm–endotherm species may have mismatched responses under climate change scenarios and the models presented here can serve as a starting point for further investigation into the biogeography of these systems. 
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  3. Abstract Background Many snakes are low-energy predators that use crypsis to ambush their prey. Most of these species feed very infrequently, are sensitive to the presence of larger vertebrates, such as humans, and spend large portions of their lifetime hidden. This makes direct observation of feeding behaviour challenging, and previous methodologies developed for documenting predation behaviours of free-ranging snakes have critical limitations. Animal-borne accelerometers have been increasingly used by ecologists to quantify activity and moment-to-moment behaviour of free ranging animals, but their application in snakes has been limited to documenting basic behavioural states (e.g., active vs. non-active). High-frequency accelerometry can provide new insight into the behaviour of this important group of predators, and here we propose a new method to quantify key aspects of the feeding behaviour of three species of viperid snakes ( Crotalus spp.) and assess the transferability of classification models across those species. Results We used open-source software to create species-specific models that classified locomotion, stillness, predatory striking, and prey swallowing with high precision, accuracy, and recall. In addition, we identified a low cost, reliable, non-invasive attachment method for accelerometry devices to be placed anteriorly on snakes, as is likely necessary for accurately classifying distinct behaviours in these species. However, species-specific models had low transferability in our cross-species comparison. Conclusions Overall, our study demonstrates the strong potential for using accelerometry to document critical feeding behaviours in snakes that are difficult to observe directly. Furthermore, we provide an ‘end-to-end’ template for identifying important behaviours involved in the foraging ecology of viperids using high-frequency accelerometry. We highlight a method of attachment of accelerometers, a technique to simulate feeding events in captivity, and a model selection procedure using biologically relevant window sizes in an open-access software for analyzing acceleration data (AcceleRater). Although we were unable to obtain a generalized model across species, if more data are incorporated from snakes across different body sizes and different contexts (i.e., moving through natural habitat), general models could potentially be developed that have higher transferability. 
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  4. Abstract Rattlesnakes are widespread mesopredators that are themselves killed and eaten by a host of other predators, including birds of prey and carnivorous mammals. Although anecdotal accounts of rattlesnake depredation are common, there are few quantitative data on encounter rates between rattlesnakes and their predators. Here we review a large database of encounters between rattlesnakes and their predators recorded from field videography of snakes in the sit-and-wait phase of their ambush hunting strategy. We found that, across 8300 hours of observation, adult rattlesnakes of six species and multiple populations exhibit low encounter rates with predators; furthermore, when predators were encountered, we never observed them to attack or kill coiled snakes. Thus, we propose that rattlesnakes are preyed upon while performing other, riskier behaviors associated with moving through the landscape. We also discuss why rattlesnakes are at low risk of predation while hunting on the surface. 
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  5. Abstract. The warming of the Arctic is affecting the carbon cycle of tundraecosystems. Most research on carbon fluxes from Arctic tundra ecosystems hasfocused on abiotic environmental controls (e.g., temperature, rainfall, orradiation). However, Arctic tundra vegetation, and therefore the carbonbalance of these ecosystems, can be substantially impacted by herbivory. Inthis study we tested how vegetation consumption by brown lemmings (Lemmus trimucronatus) canimpact carbon exchange of a wet-sedge tundra ecosystem near Utqiaġvik,Alaska during the summer and the recovery of vegetation during the followingsummer. We placed brown lemmings in individual enclosure plots and testedthe impact of lemmings' herbivory on carbon dioxide (CO2) fluxes, methane(CH4) fluxes, and the normalized difference vegetation index (NDVI)immediately after lemming removal and during the following growing season.During the first summer of the experiment, lemmings' herbivory reduced plantbiomass (as shown by the decrease in the NDVI) and decreased net CO2uptake while not significantly impacting CH4 emissions. CH4emissions were likely not significantly affected due to CH4 beingproduced deeper in the soil and escaping from the stem bases of the vascularplants. The summer following the lemming treatments, NDVI and net CO2fluxes returned to magnitudes similar to those observed before the start ofthe experiment, suggesting a complete recovery of the vegetation and atransitory nature of the impact of lemming herbivory. Overall, lemmingherbivory has short-term but substantial effects on carbon sequestration byvegetation and might contribute to the considerable interannual variabilityin CO2 fluxes from tundra ecosystems. 
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  6. Synopsis Tails are widespread in the animal world and play important roles in locomotor tasks, such as propulsion, maneuvering, stability, and manipulation of objects. Kangaroo rats, bipedal hopping rodents, use their tail for balancing during hopping, but the role of their tail during the vertical evasive escape jumps they perform when attacked by predators is yet to be determined. Because we observed kangaroo rats swinging their tails around their bodies while airborne following escape jumps, we hypothesized that kangaroo rats use their tails to not only stabilize their bodies while airborne, but also to perform aerial re-orientations. We collected video data from free-ranging desert kangaroo rats (Dipodomys deserti) performing escape jumps in response to a simulated predator attack and analyzed the rotation of their bodies and tails in the yaw plane (about the vertical-axis). Kangaroo rat escape responses were highly variable. The magnitude of body re-orientation in yaw was independent of jump height, jump distance, and aerial time. Kangaroo rats exhibited a stepwise re-orientation while airborne, in which slower turning periods corresponded with the tail center of mass being aligned close to the vertical rotation axis of the body. To examine the effect of tail motion on body re-orientation during a jump, we compared average rate of change in angular momentum. Rate of change in tail angular momentum was nearly proportional to that of the body, indicating that the tail reorients the body in the yaw plane during aerial escape leaps by kangaroo rats. Although kangaroo rats make dynamic 3D movements during their escape leaps, our data suggest that kangaroo rats use their tails to control orientation in the yaw plane. Additionally, we show that kangaroo rats rarely use their tail length at full potential in yaw, suggesting the importance of tail movement through multiple planes simultaneously. 
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  7. Abstract The outcomes of predator-prey interactions between endotherms and ectotherms can be heavily influenced by environmental temperature, owing to the difference in how body temperature affects locomotor performance. However, as elastic energy storage mechanisms can allow ectotherms to maintain high levels of performance at cooler body temperatures, detailed analyses of kinematics are necessary to fully understand how changes in temperature might alter endotherm-ectotherm predator-prey interactions. Viperid snakes are widely distributed ectothermic mesopredators that interact with endotherms both as predator and prey. Although there are numerous studies on the kinematics of viper strikes, surprisingly few have analyzed how this rapid movement is affected by temperature. Here we studied the effects of temperature on the predatory strike performance of rattlesnakes (Crotalus spp.), abundant new world vipers, using both field and captive experimental contexts. We found that the effects of temperature on predatory strike performance are limited, with warmer snakes achieving slightly higher maximum strike acceleration, but similar maximum velocity. Our results suggest that, unlike defensive strikes to predators, rattlesnakes may not attempt to maximize strike speed when attacking prey, and thus the outcomes of predatory strikes may not be heavily influenced by changes in temperature. 
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  8. ABSTRACT Movements of ectotherms are constrained by their body temperature owing to the effects of temperature on muscle physiology. As physical performance often affects the outcome of predator–prey interactions, environmental temperature can influence the ability of ectotherms to capture prey and/or defend themselves against predators. However, previous research on the kinematics of ectotherms suggests that some species may use elastic storage mechanisms when attacking or defending, thereby mitigating the effects of sub-optimal temperature. Rattlesnakes ( Crotalus spp.) are a speciose group of ectothermic viperid snakes that rely on crypsis, rattling and striking to deter predators. We examined the influence of body temperature on the behavior and kinematics of two rattlesnake species ( Crotalus oreganus helleri and Crotalus scutulatus ) when defensively striking towards a threatening stimulus. We recorded defensive strikes at body temperatures ranging from 15–35°C. We found that strike speed and speed of mouth gaping during the strike were positively correlated with temperature. We also found a marginal effect of temperature on the probability of striking, latency to strike and strike outcome. Overall, warmer snakes are more likely to strike, strike faster, open their mouth faster and reach maximum gape earlier than colder snakes. However, the effects of temperature were less than would be expected for purely muscle-driven movements. Our results suggest that, although rattlesnakes are at a greater risk of predation at colder body temperatures, their decrease in strike performance may be mitigated to some extent by employing mechanisms in addition to skeletal muscle contraction (e.g. elastic energy storage) to power strikes. 
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  9. Abstract Body size is a key factor that influences antipredator behavior. For animals that rely on jumping to escape from predators, there is a theoretical trade‐off between jump distance and acceleration as body size changes at both the inter‐ and intraspecific levels. Assuming geometric similarity, acceleration will decrease with increasing body size due to a smaller increase in muscle cross‐sectional area than body mass. Smaller animals will likely have a similar jump distance as larger animals due to their shorter limbs and faster accelerations. Therefore, in order to maintain acceleration in a jump across different body sizes, hind limbs must be disproportionately bigger for larger animals. We explored this prediction using four species of kangaroo rats (Dipodomysspp.), a genus of bipedal rodent with similar morphology across a range of body sizes (40–150 g). Kangaroo rat jump performance was measured by simulating snake strikes to free‐ranging individuals. Additionally, morphological measurements of hind limb muscles and segment lengths were obtained from thawed frozen specimens. Overall, jump acceleration was constant across body sizes and jump distance increased with increasing size. Additionally, kangaroo rat hind limb muscle mass and cross‐sectional area scaled with positive allometry. Ankle extensor tendon cross‐sectional area also scaled with positive allometry. Hind limb segment length scaled isometrically, with the exception of the metatarsals, which scaled with negative allometry. Overall, these findings support the hypothesis that kangaroo rat hind limbs are built to maintain jump acceleration rather than jump distance. Selective pressure from single‐strike predators, such as snakes and owls, likely drives this relationship. 
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