Search

Tails play essential roles in functions related to locomotor stability and maneuverability among terrestrial and arboreal animals. In kangaroo rats, bipedal hopping rodents, tails are used as effective inertial appendages for stability in hopping, but also facilitate stability and maneuverability during predator escape leaps. The complexity of tail functionality shows great potential for bio-inspiration and robotic device design, as maneuvering is accomplished by a long and light-weight inertial appendage. To (1) further understand the mechanics of how kangaroo rats use their tails during aerial maneuvers and (2) explore if we can achieve this behavior with a simplified tail-like appendage (i.e., template), we combined quantified animal observations, computational simulations, and experiments with a two degrees of freedom (2-DoF) tailed robot. We used video data from free-ranging kangaroo rats escaping from a simulated predator and analyzed body and tail motion for the airborne phase. To explain tail contributions to body orientation (i.e., spatial reorientation), we built a mid-air kangaroo rat computational model and demonstrated that the three-dimensional body orientation of the model can be controlled by a simplified 2-DoF tail with a nonlinear control strategy. Resulting simulated trajectories show movement patterns similar to those observed in kangaroo rats. Our robot experiments show that a lightweight tail can generate a large yaw displacement and stabilize pitch and roll angles to zero simultaneously. Our work contributes to better understanding of the form-function relationship of the kangaroo rat tail and lays out an important foundation for bio-inspiration in robotic devices that have lightweight tail-like appendages for mid-air maneuvering., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

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 (Dipodomys spp.), 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., (© 2021 Anatomical Society.)

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., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Human activities alter processes that control local biodiversity, causing changes in the abundance and identity of species in ecosystems. However, restoring biodiversity to a previous state is rarely as simple as reintroducing lost species or restoring processes to their pre-disturbance state. Theory suggests that established species can impede shifts in species composition via a variety of mechanisms, including direct interference, pre-empting resources or habitat alteration. These mechanisms can create transitory dynamics that delay convergence to an expected end state. We use an experimental manipulation of a desert rodent community to examine differences in recolonization dynamics of a dominant competitor (kangaroo rats of the genus Dipodomys ) when patches were already occupied by an existing rodent community relative to when patches were empty. Recovery of kangaroo rat populations was slow on plots with an established community, taking approximately 2 years, in contrast with rapid recovery on empty plots with no established residents (approx. three months). These results demonstrate that the presence of an established alternate community inhibits recolonization by new species, even those that should be dominant in the community. This has important implications for understanding how biodiversity may change in the future, and what processes may slow or prevent this change.

Background: Effective management of ground squirrels relies on an integrated pest management (IPM) approach. Rodenticides may be included in an IPM program, but they must be efficacious with minimal impact on nontarget species. A zinc phosphide-coated green bait may meet these requirements. We established a study in northeastern California to test zinc phosphide-coated cabbage as a management tool for Belding's ground squirrels (Urocitellus beldingi). We specifically addressed factors that would influence the efficacy of a baiting program, as well as potential exposure risk to nontarget species., Results: We found that prebaiting was an important application strategy, and efficacy increased as ground squirrel abundance increased. Efficacy was also greater in western portions of the study area, likely due to greater bait consumption at western sites. Belding's ground squirrels fed most heavily on cabbage during mid-morning and late afternoon; bait applications shortly before these time periods would increase bait consumption while minimizing nontarget risk. Bait uptake was greatest around burrow entrances. The only nontarget species observed feeding on cabbage was the California kangaroo rat (Dipodomys californicus), although they were never observed feeding on treated cabbage., Conclusion: Zinc phosphide-coated cabbage can be an efficacious tool for managing ground squirrels, but there will be limitations on where and how it can be used effectively. It posed a low risk to nontarget species present in our study area, but nontarget risk could vary regionally. The use of a zinc phosphide-coated green bait should only be one part of an IPM strategy for managing ground squirrels. © 2019 Society of Chemical Industry., (© 2019 Society of Chemical Industry.)

Bipedal hopping is an efficient form of locomotion, yet it remains relatively rare in the natural world. Previous research has suggested that the tail balances the angular momentum of the legs to produce steady state bipedal hopping. In this study, we employ a 3D physics simulation engine to optimize gaits for an animat whose control and morphological characteristics are subject to computational evolution, which emulates properties of natural evolution. Results indicate that the order of gene fixation during the evolutionary process influences whether a bipedal hopping or quadrupedal bounding gait emerges. Furthermore, we found that in the most effective bipedal hoppers the tail balances the angular momentum of the torso, rather than the legs as previously thought. Finally, there appears to be a specific range of tail masses, as a proportion of total body mass, wherein the most effective bipedal hoppers evolve.

Kangaroo rats are small bipedal desert rodents that use erratic vertical jumps to escape predator strikes. In this study we examined how individual hind limb joints of desert kangaroo rats ( Dipodomys deserti ) power vertical jumps across a range of heights. We hypothesized that increases in net work would be equally divided across hind limb joints with increases in jump height. To test this hypothesis, we used an inverse dynamics analysis to quantify the mechanical output from the hind limb joints of kangaroo rats jumping vertically over a wide range of heights. The kangaroo rats in this study reached maximal jump heights up to ∼9-times hip height. Net joint work increased significantly with jump height at the hip, knee and ankle, and decreased significantly at the metatarsal-phalangeal joint. The increase in net work generated by each joint was not proportional across joints but was dominated by the ankle, which ranged from contributing 56% of the work done on the center of mass at low jumps to 70% during the highest jumps. Therefore, the results of this study did not support our hypothesis. However, using an anatomical model, we estimated that a substantial proportion of the work delivered at the ankle (48%) was transferred from proximal muscles via the biarticular ankle extensors., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Many cursorial and large hopping species are extremely efficient locomotors with various morphological adaptations believed to reduce mechanical demand and improve movement efficiency, including elongated distal limb segments. However, despite having elongated limbs, small hoppers such as desert kangaroo rats ( Dipodomys deserti ) are less efficient locomotors than their larger counterparts, which may be in part due to avoiding predators through explosive jumping movements. Despite potentially conflicting mechanical demands between the two movements, kangaroo rats are both excellent jumpers and attain high hopping speeds, likely due to a specialized hindlimb musculoskeletal morphology. This study combined experimental dissection data with a static analysis of muscle moment generating capacities using a newly developed musculoskeletal model to characterize kangaroo rat hindlimb musculoskeletal architecture and investigate how morphology has evolved to meet hopping and jumping mechanical demands. Hindlimb morphology appears biased towards generating constant moment arms over large joint ranges of motion in this species, which may balance competing requirements by reducing the need for posture and movement specific excitation patterns. The ankle extensors are a major exception to the strong positive relationship exhibited by most muscles between muscle architecture parameters (e.g. L fibre ) and joint moment arms. These muscles appear suited to meeting the high moments required for jumping: the biarticular nature of the ankle extensors is leveraged to reduce MTU strain and create a four-bar linkage that facilitates proximal force transfer. The kangaroo rat hindlimb provides an interesting case study for understanding how morphology balances the sometimes competing demands of hopping and jumping., (© 2018 The Author(s).)

Predator presence causes acute stress in mammals. A prey animal's stress response increases its chance of survival during life-threatening situations through adaptive changes in behavior and physiology. Some components of the physiological stress response can lead to changes in body surface temperatures. Body temperature changes in prey could provide information about prey state to predators that sense heat, such as pit vipers. We determined whether wild rodents undergo a stress-induced change in body surface temperature upon detecting and investigating rattlesnake predators. We staged encounters between free-ranging Merriam's kangaroo rats (Dipodomys merriami) and tethered Mojave rattlesnakes (Crotalus scutulatus) at baited feeding stations, and recorded interactions with a thermal-imaging camera. Kangaroo rats showed a significant change in maximum head temperature, snout temperature, and hind leg temperature during interactions with rattlesnakes. This supports the hypothesis that presence of a predator induces body temperature changes in prey animals. If changes in prey heat signature are detectable by heat-sensitive rattlesnakes, rattlesnakes could use this information to evaluate prey vigilance or arousal before striking; however, more detailed information on the sensory ecology of the pit organ under field conditions is needed to evaluate this possibility., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Predation plays a central role in the lives of most organisms. Predators must find and subdue prey to survive and reproduce, whereas prey must avoid predators to do the same. The resultant antagonistic coevolution often leads to extreme adaptations in both parties. Few examples capture the imagination like a rapid strike from a venomous snake. However, almost nothing is known about strike performance of viperid snakes under natural conditions. We obtained high-speed (500 fps) three-dimensional video in the field (at night using infrared lights) of Mohave rattlesnakes (Crotalus scutulatus) attempting to capture Merriam's kangaroo rats (Dipodomys merriami). Strikes occurred from a range of distances (4.6 to 20.6 cm), and rattlesnake performance was highly variable. Missed capture attempts resulted from both rapid escape maneuvers and poor strike accuracy. Maximum velocity and acceleration of some rattlesnake strikes fell within the range of reported laboratory values, but some far exceeded most observations. Thus, quantifying rapid predator-prey interactions in the wild will propel our understanding of animal performance.

Population sinks present unique conservation challenges. The loss of individuals in sinks can compromise persistence; but conversely, sinks can improve viability by improving connectivity and facilitating the recolonization of vacant sources. To assess the contribution of sinks to regional population persistence of declining populations, we simulated source-sink dynamics for 3 very different endangered species: Black-capped Vireos (Vireo atricapilla) at Fort Hood, Texas, Ord's kangaroo rats (Dipodomys ordii) in Alberta, and Northern Spotted Owls (Strix occidentalis caurina) in the northwestern United States. We used empirical data from these case studies to parameterize spatially explicit individual-based models. We then used the models to quantify population abundance and persistence with and without long-term sinks. The contributions of sink habitats varied widely. Sinks were detrimental, particularly when they functioned as strong sinks with few emigrants in declining populations (e.g., Alberta's Ord's kangaroo rat) and benign in robust populations (e.g., Black-capped Vireos) when Brown-headed Cowbird (Molothrus ater) parasitism was controlled. Sinks, including ecological traps, were also crucial in delaying declines when there were few sources (e.g., in Black-capped Vireo populations with no Cowbird control). Sink contributions were also nuanced. For example, sinks that supported large, variable populations were subject to greater extinction risk (e.g., Northern Spotted Owls). In each of our case studies, new context-dependent sinks emerged, underscoring the dynamic nature of sources and sinks and the need for frequent re-assessment. Our results imply that management actions based on assumptions that sink habitats are generally harmful or helpful risk undermining conservation efforts for declining populations., (© 2015 Society for Conservation Biology.)

Species distributions are known to be limited by biotic and abiotic factors at multiple temporal and spatial scales. Species distribution models, however, frequently assume a population at equilibrium in both time and space. Studies of habitat selection have repeatedly shown the difficulty of estimating resource selection if the scale or extent of analysis is incorrect. Here, we present a multi-step approach to estimate the realized and potential distribution of the endangered giant kangaroo rat. First, we estimate the potential distribution by modeling suitability at a range-wide scale using static bioclimatic variables. We then examine annual changes in extent at a population-level. We define "available" habitat based on the total suitable potential distribution at the range-wide scale. Then, within the available habitat, model changes in population extent driven by multiple measures of resource availability. By modeling distributions for a population with robust estimates of population extent through time, and ecologically relevant predictor variables, we improved the predictive ability of SDMs, as well as revealed an unanticipated relationship between population extent and precipitation at multiple scales. At a range-wide scale, the best model indicated the giant kangaroo rat was limited to areas that received little to no precipitation in the summer months. In contrast, the best model for shorter time scales showed a positive relation with resource abundance, driven by precipitation, in the current and previous year. These results suggest that the distribution of the giant kangaroo rat was limited to the wettest parts of the drier areas within the study region. This multi-step approach reinforces the differing relationship species may have with environmental variables at different scales, provides a novel method for defining "available" habitat in habitat selection studies, and suggests a way to create distribution models at spatial and temporal scales relevant to theoretical and applied ecologists.

In this article, we first extend the superpopulation capture-recapture model to multiple states (locations or populations) for two age groups., Wen et al., (2011; 2013) developed a new approach combining capture-recapture data with population assignment information to estimate the relative contributions of in situ births and immigrants to the growth of a single study population. Here, we first generalize Wen et al., (2011; 2013) approach to a system composed of multiple study populations (multi-state) with two age groups, where an imputation approach is employed to account for the uncertainty inherent in the population assignment information. Then we develop a different, individual-level mixture model approach to integrate the individual-level population assignment information with the capture-recapture data. Our simulation and real data analyses show that the fusion of population assignment information with capture-recapture data allows us to estimate the origination-specific recruitment of new animals to the system and the dispersal process between populations within the system. Compared to a standard capture-recapture model, our new models improve the estimation of demographic parameters, including survival probability, origination-specific entry probability, and especially the probability of movement between populations, yielding higher accuracy and precision., (© 2014, The International Biometric Society.)

Identifying factors that may be responsible for regulating the size of animal populations is a cornerstone in understanding population ecology. The main factors that are thought to influence population size are either resources (bottom-up), or predation (top-down), or interspecific competition (parallel). However, there are highly variable and often contradictory results regarding their relative strengths and influence. These varied results are often interpreted as indicating "shifting control" among the three main factors, or a complex, nonlinear relationship among environmental variables, resource availability, predation, and competition. We argue here that there is a "missing link" in our understanding of predator-prey dynamics. We explore whether the landscape-of-fear model can help us clarify the inconsistencies and increase our understanding of the roles, extent, and possible interactions of top-down, bottom-up, and parallel factors on prey population abundance. We propose two main predictions derived from the landscape-of-fear model: (1) for a single species, we suggest that as the makeup of the landscape of fear changes from relatively safe to relatively risky, bottom-up impacts switch from strong to weak as top-down impacts go from weak to strong; (2) for two or more species, interspecific competitive interactions produce various combinations of bottom-up, top-down, and parallel impacts depending on the dominant competing species and whether the landscapes of fear are shared or distinctive among competing species. We contend that these predictions could successfully explain many of the complex and contradictory results of current research. We test some of these predictions based on long-term data for small mammals from the Chihuahuan Desert in the United States, and Mexico. We conclude that the landscape-of-fear model does provide reasonable explanations for many of the reported studies and should be tested further to better understand the effects of bottom-up, top-down, and parallel factors on population dynamics.

The muscle work required to sustain steady-speed locomotion depends largely upon the mechanical energy needed to redirect the centre of mass and the degree to which this energy can be stored and returned elastically. Previous studies have found that large bipedal hoppers can elastically store and return a large fraction of the energy required to hop, whereas small bipedal hoppers can only elastically store and return a relatively small fraction. Here, we consider the extent to which large and small bipedal hoppers (tammar wallabies, approx. 7 kg, and desert kangaroo rats, approx. 0.1 kg) reduce the mechanical energy needed to redirect the centre of mass by reducing collisions. We hypothesize that kangaroo rats will reduce collisions to a greater extent than wallabies since kangaroo rats cannot elastically store and return as high a fraction of the mechanical energy of hopping as wallabies. We find that kangaroo rats use a significantly smaller collision angle than wallabies by employing ground reaction force vectors that are more vertical and center of mass velocity vectors that are more horizontal and thereby reduce their mechanical cost of transport. A collision-based approach paired with tendon morphometry may reveal this effect more generally among bipedal runners and quadrupedal trotters.

Using 20 years of demographic and genetic data from four populations of banner-tailed kangaroo rats (Dipodomys spectabilis), we asked whether dispersing individuals gain benefits during adulthood that might compensate for the substantial survival costs they experience as juveniles. Compared to philopatric animals, within- and between-population dispersers gained no measureable advantages in adult survival, fecundity, or probability of recruiting offspring to adulthood. Restricting analyses to members of two central populations living more than 15 times the median dispersal distance from the study site edge, and using peripheral populations only to detect dispersal or offspring recruitment "offsite," did not change this result. Population density during year of birth had small negative effects on adult survival and fecundity, but there were no interactions with dispersal phenotype. We found no evidence that dispersers gained access to superior habitat or that their offspring suffered less inbreeding depression. Our results are consistent with fitness advantages being indirect; by leaving, dispersers release their kin from competition. Our results are also consistent with the possibility that dispersal is the "best of a bad lot." If dispersal is a conditional strategy, then the benefits may be obscured in observational data that compare dispersing individuals to philopatric individuals rather than to individuals whose dispersal phenotype is experimentally manipulated.

We hypothesize that the inner medulla of the kangaroo rat Dipodomys merriami, a desert rodent that concentrates its urine to over 6,000 mosmol/kg H(2)O, provides unique examples of architectural features necessary for production of highly concentrated urine. To investigate this architecture, inner medullary vascular segments in the outer inner medulla were assessed with immunofluorescence and digital reconstructions from tissue sections. Descending vasa recta (DVR) expressing the urea transporter UT-B and the water channel aquaporin 1 lie at the periphery of groups of collecting ducts (CDs) that coalesce in their descent through the inner medulla. Ascending vasa recta (AVR) lie inside and outside groups of CDs. DVR peel away from vascular bundles at a uniform rate as they descend the inner medulla, and feed into networks of AVR that are associated with organized clusters of CDs. These AVR form interstitial nodal spaces, with each space composed of a single CD, two AVR, and one or more ascending thin limbs or prebend segments, an architecture that may lead to solute compartmentation and fluid fluxes essential to the urine concentrating mechanism. Although we have identified several apparent differences, the tubulovascular architecture of the kangaroo rat inner medulla is remarkably similar to that of the Munich Wistar rat at the level of our analyses. More detailed studies are required for identifying interspecies functional differences.

Many models of selection predict that populations will lose variation in traits that affect fitness. Nonetheless, phenotypic variation is commonly observed in natural populations. We tested the influences of competition and spatial heterogeneity on behavioral variation within and among populations of Merriam's kangaroo rats (Dipodomys merriami) and tested for the differential expression of trait correlations. We found that populations of D. merriami exhibited more aggression at sites with more competition. Contrary to theoretical predictions and empirical results in other systems, the sites with the greatest spatial heterogeneity and highest levels of competition did not exhibit the most behavioral variation among individuals. However, the greatest within-individual behavioral variability in boldness (response to cues of predator presence) was exhibited where spatial heterogeneity was highest. Aggression and boldness of D. merriami were highly repeatable, that is, individuals behaved in a consistent manner over time, and the two behaviors were also highly correlated. Interestingly, the strength of this correlation was greatest where the competitive community was least diverse. These findings add to increasing evidence that natural populations of animals exhibit patterns of behavioral covariance, or personality structure, and suggest that competitive variation may act to erode personality structure.

1. Ecosystem engineers impact communities by altering habitat conditions, but they can also have strong effects through consumptive, competitive and other non-engineering pathways. 2. Engineering effects can lead to fundamentally different community dynamics than non-engineering effects, but the relative strengths of these interactions are seldom quantified. 3. We combined structural equation modelling and exclosure experiments to partition the effects of a keystone engineer, the giant kangaroo rat (Dipodomys ingens), on plants, invertebrates and vertebrates in a semi-arid California grassland. 4. We separated the effects of burrow creation from kangaroo rat density and found that kangaroo rats increased the diversity and abundance of other species via both engineering and non-engineering pathways. 5. Engineering was the primary factor structuring plant and small mammal communities, whereas non-engineering effects structured invertebrate communities and increased lizard abundance. 6. These results highlight the importance of the non-engineering effects of ecosystem engineers and shed new light on the multiple pathways by which strong-interactors shape communities., (© 2011 The Authors. Journal of Animal Ecology © 2011 British Ecological Society.)

Metapopulation ecology has historically been rich in theory, yet analytical approaches for inferring demographic relationships among local populations have been few. We show how reverse-time multi-state capture-recapture models can be used to estimate the importance of local recruitment and interpopulation dispersal to metapopulation growth. We use 'contribution metrics' to infer demographic connectedness among eight local populations of banner-tailed kangaroo rats, to assess their demographic closure, and to investigate sources of variation in these contributions. Using a 7 year dataset, we show that: (i) local populations are relatively independent demographically, and contributions to local population growth via dispersal within the system decline with distance; (ii) growth contributions via local survival and recruitment are greater for adults than juveniles, while contributions involving dispersal are greater for juveniles; (iii) central populations rely more on local recruitment and survival than peripheral populations; (iv) contributions involving dispersal are not clearly related to overall metapopulation density; and (v) estimated contributions from outside the system are unexpectedly large. Our analytical framework can classify metapopulations on a continuum between demographic independence and panmixia, detect hidden population growth contributions, and make inference about other population linkage forms, including rescue effects and source-sink structures. Finally, we discuss differences between demographic and genetic population linkage patterns for our system.

Translocation is used to reestablish wild populations of animals, but translocation projects often do not meet their objectives because postrelease mortality of animals is high. One reason for translocation failure is that the behavioral or ecological requirements of released animals are unmet. Maintaining founder-group social relationships during release can affect reestablishment of social species. Solitary territorial species with stable neighbors (restricted dispersal and lifetime occupation of a home range) of the same species may also benefit from the maintenance of these social relationships during translocation. We translocated Stephens' kangaroo rats (Dipodomys stephensi), a solitary species listed as endangered under the U.S. Endangered Species Act, with and without neighboring kangaroo rats. We compared the settlement (establishment of a stable home range) decisions and fitness of kangaroo rats between the 2 treatments. Kangaroo rats translocated with neighbors traveled shorter distances before establishing territories, had higher survival rates, and had significantly higher reproductive success than kangaroo rats translocated without neighbors. Number of offspring was 24-fold higher for kangaroo rats translocated with neighbors than those translocated without neighbors. Differences in behavior following release may partially explain differences in survival between the 2 groups. Immediately following release, animals translocated with neighbors fought less and spent significantly more time foraging and digging burrows than animals translocated without neighbors. Our results indicate that even for solitary species, maintaining relationships among members of a translocated group of animals can influence translocation success. This study is the first empirical demonstration of the fitness consequences of disrupting social relationships among territorial neighbors., (©2011 Society for Conservation Biology.)

Facilitation, when one species enhances the environment or performance of another species, can be highly localized in space. While facilitation in plant communities has been intensely studied, the role of facilitation in shaping animal communities is less well understood. In the Chihuahuan Desert, both kangaroo rats and harvester ants depend on the abundant seeds of annual plants. Kangaroo rats, however, are hypothesized to facilitate harvester ants through soil disturbance and selective seed predation rather than competing with them. I used a spatially explicit approach to examine whether a positive or negative interaction exists between banner-tailed kangaroo rat (Dipodomys spectabilis) mounds and rough harvester ant (Pogonomyrmex rugosus) colonies. The presence of a scale-dependent interaction between mounds and colonies was tested by comparing fitted spatial point process models with and without interspecific effects. Also, the effect of proximity to a mound on colony mortality and spatial patterns of surviving colonies was examined. The spatial pattern of kangaroo rat mounds and harvester ant colonies was consistent with a positive interspecific interaction at small scales (<10 m). Mortality risk of vulnerable, recently founded harvester ant colonies was lower when located close to a kangaroo rat mound and proximity to a mound partly predicted the spatial pattern of surviving colonies. My findings support localized facilitation of harvester ants by kangaroo rats, likely mediated through ecosystem engineering and foraging effects on plant cover and composition. The scale-dependent effect of kangaroo rats on abiotic and biotic factors appears to result in greater founding and survivorship of young colonies near mounds. These results suggest that soil disturbance and foraging by rodents can have subtle impacts on the distribution and demography of other species.

Modest increases in global temperature have been implicated in causing population extirpations and range shifts in taxa inhabiting colder environs and in ectotherms whose thermoregulation is more closely tied to environmental conditions. Many arid-adapted endotherms already experience conditions at their physiological limits, so it is conceivable that they could be similarly affected by warming temperatures. We explored how climatic variables might influence the apparent survival of the banner-tailed kangaroo rat (Dipodomys spectabilis), a rodent endemic to the Chihuahuan Desert of North America and renowned for its behavioral and physiological adaptations to arid environments. Relative variable weight, strength of variable relationships, and other criteria indicated that summer, diurnal land surface temperature (SD&#95;LST) was the primary environmental driver of apparent survival in these arid-adapted rodents. Higher temperatures had a negative effect on apparent survival, which ranged from 0.15 (SE = 0.04) for subadults to 0.50 (SE = 0.07) for adults. Elevated SD&#95;LST may negatively influence survival through multiple pathways, including increased water loss and energy expenditure that could lead to chronic stress and/or hyperthermia that could cause direct mortality. Land surface temperatures are predicted to increase by as much 6.5°C by 2099, reducing apparent survival of adults to ~0.15 in some regions of the species' range, possibly causing a shift in their distribution. The relationship between SD&#95;LST and survival suggests a mechanism whereby physiological tolerances are exceeded resulting in a reduction to individual fitness that may ultimately cause a shift in the species' range over time.

A century ago Thomas G. Lee amassed an unparalleled collection of developmental series of North American rodents such as the thirteen-lined ground squirrel, the Plains pocket gopher and Merriam's kangaroo rat. He was the first to describe the initial attachment of the squirrel blastocyst to the antimesometrial side of the uterus. The full potential of Lee's material was not realized until after his death, when it came into the possession of Mossman. The latter relied heavily on Lee's collection when writing his seminal monograph on the comparative morphogenesis of fetal membranes and much of Lee's material was subsequently described in detail by Mossman and others. It now forms part of the Harland W. Mossman Collection at the University of Wisconsin., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Megaherbivores and small burrowing mammals commonly coexist and play important functional roles in grassland ecosystems worldwide. The interactive effects of these two functional groups of herbivores in shaping the structure and function of grassland ecosystems are poorly understood. In North America's central grasslands, domestic cattle (Bos taurus) have supplanted bison (Bison bison), and now coexist with prairie dogs (Cynomys spp.), a keystone burrowing rodent. Understanding the ecological relationships between cattle and prairie dogs and their independent and interactive effects is essential to understanding the ecology and important conservation issues affecting North American grassland ecosystems. To address these needs, we established a long-term manipulative experiment that separates the independent and interactive effects of prairie dogs and cattle using a 2 x 2 factorial design. Our study is located in the Janos-Casas Grandes region of northwestern Chihuahua, Mexico, which supports one of the largest remaining complexes of black-tailed prairie dogs (C. ludovicianus). Two years of posttreatment data show nearly twofold increases in prairie dog abundance on plots grazed by cattle compared to plots without cattle. This positive effect of cattle on prairie dogs resulted in synergistic impacts when they occurred together. Vegetation height was significantly lower on the plots where both species co-occurred compared to where either or both species was absent. The treatments also significantly affected abundance and composition of other grassland animal species, including grasshoppers and banner-tailed kangaroo rats (Dipodomys spectabilis). Our results demonstrate that two different functional groups of herbivorous mammals, burrowing mammals and domestic cattle, have distinctive and synergistic impacts in shaping the structure and function of grassland ecosystems.

Mining is one of the most important industrial activities globally; however, mining processes have critical environmental impacts, as mining is a major source of metals and metalloids that contribute significantly to the pollution of soil, sediment, water and air. Heavy metals can impact the health of exposed human populations and nonhuman receptors. This study focused on arsenic because its genotoxicity is well-known. Previously, we proposed a methodology to evaluate and integrate risk from a single source affecting different biologic receptors. Here, we propose an alternative approach estimating arsenic exposure in children and kangaroo rats using probabilistic simulation with Monte Carlo modeling. The estimates are then associated to measured DNA damage and compared to both populations of children and rodents living in contaminated and in reference areas. Finally, based on the integrated analysis of the generated information, we evaluate the potential use of wild rodents (Dipodomys merriami) as a biomonitor at mining sites. Results indicate that the variation of genotoxicity in children of the reference site is approximately 2 units when compared to the children of the contaminated site. In the rodents we observed a variation of approximately 4 units between those of the reference site when compared to those living on the contaminated site. We propose that D. merriami can be used as a biomonitor organism in sites with mining activity, and that a non-lethal test can be used to evaluate risk from metal exposure.

Larval and nymphal Ixodes pacificus Cooley and Kohls, I. (Ixodes) jellisoni Cooley and Kohls, and Dermacentor occidentalis Marx were tested for host preference when simultaneously presented with a deer mouse (Peromyscus maniculatus Wagner), California kangaroo rat (Dipodomys californicus Merriam), western fence lizard (Sceloporus occidentalis Baird and Girard), and California towhee (Pipilo crissalis Vigors) in an experimental apparatus. Differences were observed in the preferences among the three species and between life stages. More larvae of all species approached and contacted hosts than did nymphs. Subadult I. pacificus entered all host-containing chambers in the highest numbers and remained on lizards most often after contact. Subadult I. jellisoni entered and remained in the chambers containing kangaroo rats, while rejecting mice, lizards, and birds as hosts. Subadult D. occidentalis most frequently entered rodent-containing chambers and contacted these hosts. After overnight exposure to all nonavian hosts, only I. pacificus parasitized and fed successfully on all three animals. I. jellisoni fed only on kangaroo rats and D. occidentalis fed only on rodents. Molting success ranged from approximately 66 to 95% among tick species and stages. We concluded that, under laboratory conditions, I. pacificus larvae and nymphs prefer western fence lizards, but also will parasitize rodents. Dermacentor occidentalis immatures use deer mice and kangaroo rats similarly, whereas I. jellisoni subadults exclusively parasitize kangaroo rats. California towhees are considerably less attractive as hosts for these three ticks. These host preferences are consistent with what is known about the natural feeding habits of all three ticks.

Using long-term data on two kangaroo rats in the Chihuahuan Desert of North America, we fitted logistic models including the exogenous effects of seasonal rainfall patterns. Our aim was to test the effects of intraspecific interactions and seasonal rainfall in explaining and predicting the numerical fluctuations of these two kangaroo rats. We found that logistic models fit both data sets quite well; Dipodomys merriami showed lower maximum per capita growth rates than Dipodomys ordii, and in both cases logistic models were nonlinear. Summer rainfall appears to be the most important exogenous effect for both rodent populations; models including this variable were able to predict independent data better than models including winter rainfall. D. merriami was also negatively affected by another kangaroo rat (Dipodomys spectabilis), consistent with previous experimental evidence. We hypothesized that summer rainfall influences the carrying capacity of the environment by affecting seed availability and the intensity of intraspecific competition.

Behavioural syndromes, correlations of behaviours conceptually analogous to personalities, have been a topic of recent attention due to their potential to explain trade-offs in behavioural responses, apparently maladaptive behaviour and limits to plasticity. Using Merriam's kangaroo rats (Dipodomys merriami), we assessed the explanatory power and generality of hypothesized syndrome structures derived from the literature and the natural history of the species. Several aspects of functionally distinct behavioural responses of D. merriami were quantified. Syndrome structures were compared using structural equation modelling and model selection procedures. A domain-general behavioural syndrome incorporating cross-functional relationships between measures of boldness, agonistic behaviour, flexibility and food hoarding best explained the data. This pattern suggests that D. merriami behaviours should not be viewed as discrete elements but as components of a multivariate landscape. Our results support arguments that a lack of independence between behaviours may be a general aspect of behavioural phenotypes and suggest that the ability of D. merriami's behaviour to respond to selection may be constrained by underlying connections.

Locating birthplaces using genetic parentage determination can increase the precision and accuracy with which animal dispersal patterns are established. We re-analyse patterns of movement away from the birthplace as a function of time, sex and population density for a sample of 303 banner-tailed kangaroo rats, Dipodomys spectabilis. We located birth sites using a combination of likelihood-based parentage analysis with live-trapping of mothers during the breeding season. The results demonstrate that natal-breeding site distances are density dependent in this species; in particular, both sexes emigrate earlier in the year, and females disperse farther than males, at low population densities. Banner-tailed kangaroo rats were chosen as a study system because live-trapping easily detects maternal and offspring locations; nevertheless, parentage analysis reveals that some offspring evade early detection and move substantial distances before their first capture. In a few cases, the approach even detects dispersal out of the natal 'deme' prior to first capture. Parentage analysis confirms the extreme philopatry of both sexes but indicates that prior estimates of median dispersal distance were too low. For D. spectabilis, more accurate location of individual birthplaces clarifies patterns of sex bias and density dependence in dispersal, and may resolve apparent discrepancies between direct and indirect estimates of dispersal distance. For species in which mothers can be more reliably trapped than juveniles, using offspring genotypes to locate parents is a novel way that genetic techniques can contribute to the analysis of animal dispersal.

Knowledge of animal energetics is based largely upon indirect calorimetry, which is estimation of metabolic heat production by an organism from measurement of indices such as oxygen consumption or carbon dioxide production. Remarkably, indirect calorimetry has been validated by comparison to direct measurements of metabolic heat production (direct calorimetry) only for highly restricted conditions, primarily with a few species of medium-to-large mammals. Taxa with differing physiologies are little studied. For birds, for example, validations are limited to chickens and waterfowl exposed to mild environmental conditions and typically fasted for prolonged periods. Workers rely upon these restricted validations when studying animals ranging greatly in activity, phylogeny, body size and nutritional status. We tested the accuracy of respirometric estimates of energy expenditure by simultaneous indirect and direct calorimetry in a small mammal (the kangaroo rat Dipodomys merriami Mearns), a small bird (the dove Columbina inca Lesson) and a medium-sized bird (the quail Coturnix communis Linnaeus). We find that conventional respirometric estimates of energy expenditure may incorporate large errors (up to 38%) that are sufficient to call into question generalizations regarding patterns of animal energy use in many studies.

Low-frequency distortion-product otoacoustic emissions were measured in two species of kangaroo rats to test the prediction that a large footdrumming species (e.g., Dipodomys spectabilis) would have greater distortion-product otoacoustic emission amplitude than a small non-footdrumming species (e.g., Dipodomys merriami), indicating better hearing sensitivity at low frequencies. Equal-level (65 dB SPL) stimulus tones ( f(1), f(2)), presented over a ( f(1)) range of 200-1000 Hz, were used to evoke the 2 f(1)- f(2) distortion-product otoacoustic emission. Mean 2 f(1)- f(2) levels for D. merriami showed good correspondence to previously published audiograms for that species. Mean 2 f(1)- f(2) levels and 95% confidence intervals indicated species differences below 400 Hz, supporting the theory that low-frequency hearing sensitivity is better in large kangaroo rat species. These results suggest that the size-related divergence in footdrumming behavior may be related to differential auditory sensitivity.

Extensive mark-recapture data from banner-tailed kangaroo rats, Dipodomys spectabilis, have shown that both males and females are highly philopatric and suggest the possibility of close inbreeding. However, indirect analyses based on genetic structure appear to contradict direct observations, suggesting longer dispersal distances. Using microsatellite genotypes from most members of a banner-tailed kangaroo rat population during five successive breeding seasons, we ask how relatedness is influenced by dispersal and how it in turn influences mating patterns. The data confirm that, because of philopatry, neighbours are often close relatives. However, patterns of parentage also show that the average distance between mates is large relative to natal dispersal distances and larger than the average distance between nearest opposite-sexed neighbours. Females' mates were often not their nearest male neighbour and many were less related than the nearest male neighbour. We detected multiple paternity in some females' litters; both sexes produce offspring with multiple mates within and between breeding seasons. At the population level, heterozygosities were high and estimates of F were low, indicating that levels of inbreeding were low. Using individual inbreeding coefficients of all juveniles to estimate their parents' relatedness, we found that parental relatedness was significantly lower than relatedness between nearest opposite-sexed adult neighbours. Thus in philopatric populations, long breeding forays can cause genes to move further than individuals disperse, and polyandry may serve to reduce relatedness between mates.

Desert endotherms such as Merriam's kangaroo rat (Dipodomys merriami) use both behavioral and physiological means to conserve energy and water. The energy and water needs of kangaroo rats are affected by their thermal environment. Animals that choose temperatures within their thermoneutral zone (TNZ) minimize energy expenditure but may impair water balance because the ratio of water loss to water gain is high. At temperatures below the TNZ, water balance may be improved because animals generate more oxidative water and reduce evaporative water loss; however, they must also increase energy expenditure to maintain a normal body temperature. Hence, it is not possible for kangaroo rats to choose thermal environments that simultaneously minimize energy expenditure and increase water conservation. I used a thermal gradient to test whether water stress, energy stress, simultaneous water and energy stress, or no water/energy stress affected the thermal environment selected by D. merriami. During the night (i.e., active phase), animals in all four treatments chose temperatures near the bottom of their TNZ. During the day (i.e., inactive phase), animals in all four treatments settled at temperatures near the top of their TNZ. Thus, kangaroo rats chose thermal environments that minimized energy requirements, not water requirements. Because kangaroo rats have evolved high water use efficiency, energy conservation may be more important than water conservation to the fitness of extant kangaroo rats.

Desert-dwelling mammals such as Merriam's kangaroo rat (Dipodomys merriani) need to conserve both energy and water to survive desert conditions characterized by aridity and low productivity. The thyroid hormone thyroxine increases both basal metabolic rate and urinary water loss in mammals. Increases in basal metabolism and urinary water loss are likely to be detrimental to D. merriami, therefore the regulation of this hormone may be important. To examine the effects of thyroxine in this species, we implanted adult kangaroo rats with pellets designed to release specific doses of thyroxine at a constant rate for 90 days or a placebo pellet. We measured plasma thyroxine concentration, basal metabolic rate, food consumption, urine concentration and water loss in all implanted animals. Thyroxine implants significantly increased both plasma thyroxine and basal metabolic rate in a relatively dose-dependent manner. In response to thyroxine. kangaroo rats increased food consumption only slightly, but this small increase was sufficient to compensate for their elevated metabolic rates. Neither urine concentration nor water loss varied among treatment groups. Thyroxine increased energy expenditure but not water loss in this species.

Understanding the evolution of physiological traits requires considering three nonexclusive mechanisms that underlie phenotypes and cause their change over different time scales: acclimation, developmental plasticity, and natural selection for genetically fixed traits. Physiological adjustments to changes in the desiccating potential of the environment were investigated with one subspecies of common desert rodent, Dipodomys merriami merriami (Merriam's kangaroo rat). We raised young whose parents originated from environments that differ in both temperature and humidity. These young were raised under either desiccating or water-abundant conditions, and their water loss was measured at a series of temperatures to determine the effect developmental conditions have on resistance to desiccation. We then determined the contribution of acclimation to desiccation resistance by keeping the differentially raised young in conditions opposite to those during their development and again measuring water loss. We found that developmental plasticity and acclimation can completely account for the existing intraspecific variability in desiccation resistance under certain conditions. In fact, developmental and acclimatory changes can equal genetically based differences of the populations. This phenotypic plasticity can operate relatively quickly and therefore may attenuate the actions of natural selection. Understanding the extent and nature of such flexibility is critical to our understanding intraspecific variability and the consequences of changing climate.

Because individual species can play key roles, the loss of species through extinction or their gain through colonization can cause major changes in ecosystems. For almost 20 years after kangaroo rats were experimentally removed from a Chihuahuan desert ecosystem in the United States, other rodent species were unable to compensate and use the available resources. This changed abruptly in 1995, when an alien species of pocket mouse colonized the ecosystem, used most of the available resources, and compensated almost completely for the missing kangaroo rats. These results demonstrate the importance of individual species and of colonization and extinction events in the structure and dynamics of ecosystems.

A long-held assertion has been that nocturnality is an escape mechanism for many nocturnal desert rodents because of limited tolerances to heat. To test this claim, we used a treadmill to examine the tolerances to high ambient temperatures (T(a)'s) of one subspecies of desert rodent, Merriam's kangaroo rat, Dipodomys merriami merriami, from contrasting environments. We simultaneously measured body temperature (T(b)), evaporative water loss, and metabolic rates at an ecologically relevant speed (0.6 km h(-1)) at different ambient temperatures (Ta=25 degrees -42.5 degrees C). We hypothesized that kangaroo rats from a more xeric site would have greater abilities to remain active and maintain stable T(b) than those from a more mesic site, but mesic- and xeric-site animals had comparable tolerances and were active until Tb=42 degrees C. At Ta=42.5 degrees C, however, T(b) of mesic-site animals increased more quickly than in xeric-site animals. Although most animals could not run more than 18 min at Ta=42.5 degrees C, most could run at Ta=40 degrees C for at least 30 min. Benefits of nocturnality for this species may reside more in purposes of water conservation and avoidance of predation and less on the direct regulation of T(b), as T(b) is more labile than commonly thought.

Previous estimates suggested that ventilatory evaporation constitutes the major source of water loss in kangaroo rats (Dipodomys spp.). We quantified rates of water loss in Merriam's kangaroo rat (Dipodomys merriami) and demonstrate the degree to which acclimation to a particular thermal and hydric environment plays a role in the intraspecific variation in water loss evident in this species. We draw the following conclusions: (1) that water loss varies intraspecifically in Merriam's kangaroo rat, in association with habitats of contrasting aridity and temperature; (2) that animals from more xeric locations have lower water loss rates than those from more mesic sites; (3) that most water loss is cutaneous, with ventilatory evaporative water loss contributing, at most, only 44% to total evaporative water loss; and (4) that intraspecific differences in rates of water loss are not acclimatory, but fixed. After acclimating under the same conditions, xeric-site animals still show a 33% lower rate of evaporative water loss than mesic-site animals.

Banner-tailed kangaroo rats, Dipodomys spectabilis, footdrum to produce substrate-borne and airborne acoustic energy. Previous studies show that they communicate territorial ownership via airborne footdrumming signals. The research reported here used simulated footdrum patterns generated by an artificial 'thumper' to address the question of whether kangaroo rats communicate through seismic components of these acoustic signals. With microphones suspended in sealed burrows, we found that airborne sounds were attenuated by approximately 40 dB as they passed through the burrow wall into the burrow chamber. The substrate-borne vibrations from the thumper yielded sound approximately 40 dB greater in peak amplitude than the attenuated airborne sound. Thus, 99.9% of the peak power of the thumper was transmitted directly through the substrate into the burrow. The rats in sealed burrows timed their responses to playbacks of footdrums from the thumper and a loudspeaker so they did not initiate a drumming sequence during either the seismic or airborne signals. When these signals were masked by loud noise, the rats continued to drum to the seismic signal but drummed randomly during the airborne playback. These results suggest that the sealed burrow provides a quiet place in which D. spectabilis can listen for substrate-borne communications from conspecifics.

The aim of this study was to compare the contractile and series elastic properties of terrestrial mammals that use bipedal versus quadrupedal gaits. The gastrocnemius muscle of the hopping mouse (body mass 30.2 +/- 2.4 g, mean +/- S.D.) and the rat (313 +/- 10.7 g) were compared with data from the literature for the wallaby and the kangaroo rat to distinguish scaling effects and locomotion-related effects on muscle properties. Contractile length-force properties and series elastic stiffness were measured in situ during maximal tetanic contractions. The rat had a larger muscle-fibre-to-tendon-length ratio. The rat and hopping mouse showed similar normalised length-force characteristics of the gastrocnemius. Normalised stiffness in the hopping mouse was higher. The hopping mouse showed a higher capacity to store elastic energy per unit of contractile work capacity, as well as per unit of body mass. Accounting for body size differences, the rat had a smaller relative muscle mass and thus smaller work capacity than the three hopping animals considered. This is an agreement with a quadrupedal versus bipedal locomotion style. The differences in contractile and elastic properties of the gastrocnemius of the rat and hopping mouse seem to be closely related to locomotion patterns. Small animals seem to be able to utilise the storage and release of elastic energy to a far lesser extent than larger animals. However, even in animals as small as hopping mice, the storage and utilisation of elastic energy during locomotion is of functional significance and probably depends on locomotor behaviour.

Analyses of long-term experimental data from the Chihuahuan desert revealed that species diversity of other rodents was higher on plots from which kangaroo rats (Dipodomys spp.) had been removed. The difference was due to consistently higher colonization and lower extinction probabilities of small granivorous rodents in the absence of competitively dominant kangaroo rats. The results of this ecosystem experiment demonstrate the importance of both competitive exclusion and metapopulation dynamics for biological diversity in a natural community.

Two species of wild rodents (Ord's kangaroo rat, Dipodomys ordii Woodhouse, and the white-footed mouse, Peromyscus leucopus Rafinesque) were fed 0, 2, and 20% (AI) by weight concentrations of carbaryl-bran grasshopper bait in the laboratory to determine whether the bait would be consumed, the total quantity of bait consumed within a specified period, whether mortality would occur, and whether bait would be consumed if other foods were readily available. Consumption tests showed that bait ingestion was inversely proportional to concentration. No mortalities occurred at the 0% level, whereas the 2 and 20% concentration treatments resulted in 18.8 and 97.0% mortality, respectively. The apparent repellency of 20% bait indicated that starvation may have caused the deaths. Food preference tests disclosed that D. ordii and P. leucopus preferred natural forage rather than treated and untreated wheat bran in the laboratory.

Previous studies have demonstrated gonadal control of mammalian scent glands; castration leads to reduced scent-marking rates and smaller gland sizes. I investigated whether gonadal hormones control the size of the dorsal gland (a specialized sebaceous gland) and sandbathing as a scent-marking behavior in adult male and female bannertail kangaroo rats (Dipodomys spectabilis). Gland sizes of males and females were similar in all age classes, except adult males with larger body weights had proportionately larger glands than females. Male gland sizes declined 18% following castration but were not significantly smaller than those of intact males; females showed no change in dorsal gland size either as a result of ovariectomy or after estradiol benzoate implants. Sandbathing rates also did not decline as a result of gonadectomy. Rather, castrated males and overiectomized females sandbathed at higher frequencies than intact males and ovariectomized females with estradiol benzoate implants, respectively. Gonadal hormones apparently have little influence on the function of a specialized scent gland and may inhibit sandbathing as a scent-marking behavior in adult D. spectabilis.