Cranial variation in species and subspecies of kangaroo rats (Dipodomys, Dipodomyinae, Rodentia) according to geometric morphometrics.

Traditional Dipodomys (sub)species identification uses geography, phenotype, and external/skull measurements. Such measurements are correlated with size and thus redundant. I assessed the value of scaled cranial shape, based on two-dimensional landmarks (analyzed using geometric morphometric methods) in distinguishing Dipodomys taxa, and in summarizing their variation. My dataset includes 601 adult specimens from 20 species (49 operational taxonomic units - OTUs) across 190 localities. Cranial shape was highly useful in classifying Dipodomys taxa without considering geography. The auditory bulla was the most variable region-taxa differed in its hypertrophy, accompanied by different degrees of nearby structure crowding. Cranial shape was weakly allometric, with no significant sexual dimorphism. Weak size dimorphism was detected. (Sub)specific taxonomy is not reflective of shape variation, as the number of subspecies per species is not associated with disparity. Shape had significant phylogenetic signal, but subspecies did not always cluster with conspecifics and species did not always cluster according to phylogenetic relationship/taxonomy. Shape variation was correlated with climate, and species differed in morphological disparity and degree of specialization, which may contribute to divergence in shape variation patterns from phylogeny. D. deserti was the most specialized species, diverging greatly from the genus mean; D. heermanni was the least specialized. This study provides new insights into morphological variation of North American keystone species, several of conservation interest, for example, D. heermanni berkeleyensis, D. h. dixoni, D. nitratoides brevinasus, and D. n. nitratoides.

Ecological factors shaping ectoparasite communities on heteromyid rodents at Médanos de Samalayuca.

Rodent ectoparasites are vectors for important pathogens of wildlife, domestic animals, and even zoonosis. Nevertheless, distribution patterns of ectoparasites are not fully understood; habitat, season, and host species are important predictors of distribution and prevalence. Heteromyid rodents are considered important reservoirs of diseases, given the presence of different ectoparasites and pathogens in them, and they offer the opportunity to learn about the ecology of parasites. The aim of the present work was to survey ectoparasites associated with heteromyid rodents near a National Protected Area in Chihuahua Mexico, south of the USA-Mexico border, and asses the effects of ecological factors (season, vegetation type, host species, and host body condition) on parasite infestation. We sampled five different locations from January 2018 to July 2022; 845 heteromyid rodents were examined and 49 fleas and 33 ticks were collected. Ectoparasites belonged to the Siphonaptera and Ixodida orders, including three families Ixodidae (Riphicephalus sanguineus), Pulicidae (Pulex irritans), and Ctenophthalmidae (Meringins altipecten, M. dipodomys). Five species of host rodents were captured, Dipodomys merriami, D. ordii, Chaetodipus eremicus, C. hispidus, and C. intermedius, but the last two species did not present any ectoparasites. Dipodomys merriami presented the highest flea and tick prevalence followed by D. ordii. We found parasitic partnerships between heteromyids according to ecological factors. The infestation in C. eremicus was related to body condition, vegetation type, and sex; in D. merriami, it was related to vegetation type and season, while D. ordii did not present a clear pattern of infestation. Our results suggest that the infestation patterns of heteromyid rodents in desert habitats are species dependent.

Incomplete recovery of seeds from scatterhoards by granivorous rodents: Implications for plant establishment.

Granivorous rodents are important components of ecosystems not only because they consume seeds but also because some aid in seed dispersal through seed-caching behaviors. Some rodents bury seeds in shallow pits throughout territories, called scatterhoards, that individuals recover, pilfer, or transfer to other caches. We suspect some single-seed caches in environments represent missed seeds from reclaiming or pilfering caches. We documented the sloppiness of seed removal from scatterhoards of soapweed yucca (Yucca glauca) seeds by Ord's kangaroo rats (Dipodomys ordii). We quantified the frequency and location of seeds remaining. In an experiment with artificial caches of three sizes, kangaroo rats harvested 51% of caches after one night, and 53% had incomplete recovery with at least one seed remaining. The greater the number of seeds in caches, the greater frequency of incomplete recovery. In another experiment with natural and artificial caches, 75% of caches were excavated after 8 days, with at least 70% having at least one seed remaining. Regardless of original cache size, a single seed represented the mode for seeds remaining. Incomplete recovery of seeds likely benefits plant establishment, potentially significantly in some systems. Remaining seeds, especially those buried at bottoms of caches, likely will stay undetected in landscapes, yielding propagules for subsequent plant generations. Soapweed yucca has large but light, flat wind-dispersed seeds, and removal of caches with smaller seeds might have greater frequency of missed seeds during recovery and pilfering by rodents. Our results suggest that scatter-hoarding granivores also contribute to plant establishment by leaving limited numbers of seeds behind when removing caches, at least in some systems.

Pilfering of western juniper seed caches by scatter-hoarding rodents varies by microsite and canopy type.

Scatter-hoarding rodents store seeds throughout their home ranges in superficially buried caches which, unlike seeds larder-hoarded in burrows, are difficult to defend. Cached seeds are often pilfered by other scatter-hoarders and either re-cached, eaten or larder-hoarded. Such seed movements can influence seedling recruitment, because only seeds remaining in caches are likely to germinate. Although the importance of scatter-hoarding rodents in the dispersal of western juniper seeds has recently been revealed, the level of pilfering that occurs after initial burial is unknown. Seed traits, soil moisture, and substrate can influence pilfering processes, but less is known about how pilfering varies among caches placed in open versus canopy microsites, or how cache discovery and removal varies among different canopy-types, tree versus shrub. We compared the removal of artificial caches between open and canopy microsites and between tree and shrub canopies at two sites in northeastern California during late spring and fall. We also used trail cameras at one site to monitor artificial cache removal, identify potential pilferers, and illuminate microsite use by scatter-hoarders. Removal of artificial caches was faster in open microsites at both sites during both seasons, and more caches were removed from shrub than tree canopies. California kangaroo rats were the species observed most on cameras, foraging most often in open microsites, which could explain the observed pilfering patterns. This is the first study to document pilfering of western juniper seeds, providing further evidence of the importance of scatter-hoarding rodent foraging behavior in understanding seedling recruitment processes in juniper woodlands.

Does urbanization ameliorate the effect of endoparasite infection in kangaroo rats?

Urban development can fragment and degrade remnant habitat. Such habitat alterations can have profound impacts on wildlife, including effects on population density, parasite infection status, parasite prevalence, and body condition. We investigated the influence of urbanization on populations of Merriam's kangaroo rat (Dipodomys merriami) and their parasites. We predicted that urban development would lead to reduced abundance, increased parasite prevalence in urban populations, increased probability of parasite infection for individual animals, and decreased body condition of kangaroo rats in urban versus wildland areas. We live trapped kangaroo rats at 5 urban and 5 wildland sites in and around Las Cruces, NM, USA from 2013 to 2015, collected fecal samples from 209 kangaroo rats, and detected endoparasites using fecal flotation and molecular barcoding. Seven parasite species were detected, although only two parasitic worms, Mastophorus dipodomis and Pterygodermatites dipodomis, occurred frequently enough to allow for statistical analysis. We found no effects of urbanization on population density or probability of parasite infection. However, wildland animals infected with P. dipodomis had lower body condition scores than infected animals in urban areas or uninfected animals in either habitat. Our results suggest that urban environments may buffer Merriam's kangaroo rats from the detrimental impacts to body condition that P. dipodomis infections can cause.

Competition shapes individual foraging and survival in a desert rodent ensemble.

Intraspecific variation, including individual diet variation, can structure populations and communities, but the causes and consequences of individual foraging strategies are often unclear. Interactions between competition and resources are thought to dictate foraging strategies (e.g. specialization vs. generalization), but classical paradigms such as optimal foraging and niche theory offer contrasting predictions for individual consumers. Furthermore, both paradigms assume that individual foraging strategies maximize fitness, yet this prediction is rarely tested. We used repeated stable isotope measurements (δ13 C, δ15 N; N = 3,509) and 6 years of capture-mark-recapture data to quantify the relationship between environmental variation, individual foraging and consumer fitness among four species of desert rodents. We tested the relative effects of intraspecific competition, interspecific competition, resource abundance and resource diversity on the foraging strategies of 349 individual animals, and then quantified apparent survival as function of individual foraging strategies. Consistent with niche theory, individuals contracted their trophic niches and increased foraging specialization in response to both intraspecific and interspecific competition, but this effect was offset by resource availability and individuals generalized when plant biomass was high. Nevertheless, individual specialists obtained no apparent fitness benefit from trophic niche contractions as the most specialized individuals exhibited a 10% reduction in monthly survival compared to the most generalized individuals. Ultimately, this resulted in annual survival probabilities nearly 4× higher for generalists compared to specialists. These results indicate that competition is the proximate driver of individual foraging strategies, and that diet-mediated fitness variation regulates population and community dynamics in stochastic resource environments. Furthermore, our findings show dietary generalism is a fitness maximizing strategy, suggesting that plastic foraging strategies may play a key role in species' ability to cope with environmental change.

Declines in rodent abundance and diversity track regional climate variability in North American drylands.

Regional long-term monitoring can enhance the detection of biodiversity declines associated with climate change, improving future projections by reducing reliance on space-for-time substitution and increasing scalability. Rodents are diverse and important consumers in drylands, regions defined by the scarcity of water that cover 45% of Earth's land surface and face increasingly drier and more variable climates. We analyzed abundance data for 22 rodent species across grassland, shrubland, ecotone, and woodland ecosystems in the southwestern USA. Two time series (1995-2006 and 2004-2013) coincided with phases of the Pacific Decadal Oscillation (PDO), which influences drought in southwestern North America. Regionally, rodent species diversity declined 20%-35%, with greater losses during the later time period. Abundance also declined regionally, but only during 2004-2013, with losses of 5% of animals captured. During the first time series (wetter climate), plant productivity outranked climate variables as the best regional predictor of rodent abundance for 70% of taxa, whereas during the second period (drier climate), climate best explained variation in abundance for 60% of taxa. Temporal dynamics in diversity and abundance differed spatially among ecosystems, with the largest declines in woodlands and shrublands of central New Mexico and Colorado. Which species were winners or losers under increasing drought and amplified interannual variability in drought depended on ecosystem type and the phase of the PDO. Fewer taxa were significant winners (18%) than losers (30%) under drought, but the identities of winners and losers differed among ecosystems for 70% of taxa. Our results suggest that the sensitivities of rodent species to climate contributed to regional declines in diversity and abundance during 1995-2013. Whether these changes portend future declines in drought-sensitive consumers in the southwestern USA will depend on the climate during the next major PDO cycle.

Rodents' responses to manipulated plant litter and seed densities: implications for restoration.

Rodent populations in arid grasslands do not always track seed production, possibly due to high levels of plant litter. When natural disturbances are suppressed, litter accumulates becoming physically complex, causing rodents to harvest fewer seeds per equivalent time foraging. It also alters security from predation. Restoring natural disturbances may be an important element in conserving rodent communities. The aim of this study was to assess the influence of two levels of plant litter cover and seed densities on nocturnal rodent population characteristics in a semiarid grassland. Specifically, I hypothesized that kangaroo rats, pocket mice, grasshopper mice, and total rodents would be higher in the sparse plant litter treatment than dense litter, whereas deer mice would be lower in sparse plots. I further hypothesized that kangaroo rats and deer mice would be higher in the seed augmented treatment compared to the unseeded treatment. A prescribed fire removed litter in four of eight plots prior to sowing native seeds 1 year postfire into two burned and two unburned plots. Rodents were live-trapped during spring and fall 1 year. Sparse litter treatment had higher total rodent abundance, biomass, and frequency of offspring compared to dense plots indicating use of stored seeds. Banner-tailed kangaroo rats had higher abundance, implying reduced predation risk. Pocket mice body mass was greater in dense plots. After winter, seeded plots had higher kangaroo rat body mass and grasshopper mice abundance than unseeded, reflecting the use of stored seeds. These short term results demonstrate litter's physical complexity may be equivalent to seed pulses on the responses of nocturnal rodents. Managers might positively influence grassland rodents by providing a mosaic of varying levels of plant litter.

Established rodent community delays recovery of dominant competitor following experimental disturbance.

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.

Historical Population Size Change and Differentiation of Relict Populations of the Endangered Giant Kangaroo Rat.

From a conservation management perspective it is important to understand how genetic diversity is partitioned across a species' range, including 1) identification of evolutionarily distinct units versus those recently isolated through anthropogenic activities and 2) the relative genetic contributions among components of fragmented (meta)populations. To address these questions, we investigated the phylogeography and metapopulation structure among relict populations of the endangered giant kangaroo rat (Dipodomys ingens) in the highly altered San Joaquin Desert Ecosystem. This keystone species underwent a ~97% range reduction over the past century, resulting in a current range that is highly fragmented, with 2 dominant northern and southern populations occurring 150 km apart. We sequenced >800 bp of mitochondrial DNA and genotyped 17 nuclear microsatellites in >275 D. ingens to assess the evolutionary relationship of these populations as well as the genetic structure within the northern metapopulation. A Bayesian Skyline Plot indicated that the species experienced a demographic expansion toward the end of the Pleistocene, with a recent population decline. Northern and southern D. ingens split 1857-13 443 years ago, prior to the massive conversion of the San Joaquin Valley to irrigated agriculture. We recommend that the northern and southern populations of D. ingens be re-classified as distinct population segments under the United States Endangered Species Act. We also observed population structure and asymmetrical migration within northern D. ingens where the Tumey Hills acted as a source contributing gene flow to all peripheral populations. This emphasized the importance of this location in the conservation of the metapopulation as a whole.

Functional connectivity and home range inferred at a microgeographic landscape genetics scale in a desert-dwelling rodent.

Gene flow in animals is limited or facilitated by different features within the landscape matrix they inhabit. The landscape representation in landscape genetics (LG) is traditionally modeled as resistance surfaces (RS), where novel optimization approaches are needed for assigning resistance values that adequately avoid subjectivity. Also, desert ecosystems and mammals are scarcely represented in LG studies. We addressed these issues by evaluating, at a microgeographic scale, the effect of landscape features on functional connectivity of the desert-dwelling Dipodomys merriami. We characterized genetic diversity and structure with microsatellites loci, estimated home ranges and movement of individuals using telemetry-one of the first with rodents, generated a set of individual and composite environmental surfaces based on hypotheses of variables influencing movement, and assessed how these variables relate to individual-based gene flow. Genetic diversity and structure results evidenced a family-induced pattern driven by first-order-related individuals, notably determining landscape genetic inferences. The vegetation cover and soil resistance optimized surface (NDVI) were the best-supported model and a significant predictor of individual genetic distance, followed by humidity and NDVI+humidity. Based on an accurate definition of thematic resolution, we also showed that vegetation is better represented as continuously (vs. categorically) distributed. Hence, with a nonsubjective optimization framework for RS and telemetry, we were able to describe that vegetation cover, soil texture, and climatic variables influence D. merriami's functional connectivity at a microgeographic scale, patterns we could further explain based on the home range, habitat use, and activity observed between sexes. We describe the relationship between environmental features and some aspects of D. merriami's behavior and physiology.

Jumping mechanics of desert kangaroo rats.

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.

Precipitation alters interactions in a grassland ecological community.

Climate change is transforming precipitation regimes world-wide. Changes in precipitation regimes are known to have powerful effects on plant productivity, but the consequences of these shifts for the dynamics of ecological communities are poorly understood. This knowledge gap hinders our ability to anticipate and mitigate the impacts of climate change on biodiversity. Precipitation may affect fauna through direct effects on physiology, behaviour or demography, through plant-mediated indirect effects, or by modifying interactions among species. In this paper, we examined the response of a semi-arid ecological community to a fivefold change in precipitation over 7 years. We examined the effects of precipitation on the dynamics of a grassland ecosystem in central California from 2007 to 2013. We conducted vegetation surveys, pitfall trapping of invertebrates, visual surveys of lizards and capture-mark-recapture surveys of rodents on 30 plots each year. We used structural equation modelling to evaluate the direct, indirect and modifying effects of precipitation on plants, ants, beetles, orthopterans, kangaroo rats, ground squirrels and lizards. We found pervasive effects of precipitation on the ecological community. Although precipitation increased plant biomass, direct effects on fauna were often stronger than plant-mediated effects. In addition, precipitation altered the sign or strength of consumer-resource and facilitative interactions among the faunal community such that negative or neutral interactions became positive or vice versa with increasing precipitation. These findings indicate that precipitation influences ecological communities in multiple ways beyond its recognized effects on primary productivity. Stochastic variation in precipitation may weaken the average strength of biotic interactions over time, thereby increasing ecosystem stability and resilience to climate change.

A mast-seeding desert shrub regulates population dynamics and behavior of its heteromyid dispersers.

Granivorous rodent populations in deserts are primarily regulated through precipitation-driven resource pulses rather than pulses associated with mast-seeding, a pattern more common in mesic habitats. We studied heteromyid responses to mast-seeding in the desert shrub blackbrush (Coleogyne ramosissima), a regionally dominant species in the Mojave-Great Basin Desert transition zone. In a 5-year study at Arches National Park, Utah, USA, we quantified spatiotemporal variation in seed resources in mast and intermast years in blackbrush-dominated and mixed desert vegetation and measured responses of Dipodomys ordii (Ord's kangaroo rat) and Perognathus flavescens (plains pocket mouse). In blackbrush-dominated vegetation, blackbrush seeds comprised >79% of seed production in a mast year, but 0% in the first postmast year. Kangaroo rat abundance in blackbrush-dominated vegetation was highest in the mast year, declined sharply at the end of the first postmast summer, and then remained at low levels for 3 years. Pocket mouse abundance was not as strongly associated with blackbrush seed production. In mixed desert vegetation, kangaroo rat abundance was higher and more uniform through time. Kangaroo rats excluded the smaller pocket mice from resource-rich patches including a pipeline disturbance and also moved their home range centers closer to this disturbance in a year of low blackbrush seed production. Home range size for kangaroo rats was unrelated to seed resource density in the mast year, but resource-poor home ranges were larger (P < 0.001) in the first postmast year, when resources were limiting. Blackbrush seeds are higher in protein and fat but lower in carbohydrates than the more highly preferred seeds of Indian ricegrass (Achnatherum hymenoides) and have similar energy value per unit of handling time. Kangaroo rats cached seeds of these two species in similar spatial configurations, implying that they were equally valued as stored food resources. Blackbrush mast is a key resource regulating populations of kangaroo rats in this ecosystem.

The complete mitochondrial genome of the Dipodomys ordii (Ord's kangaroo rat).

Ord's kangaroo rat is a kangaroo rat native to western North America. In this study, we first reported the complete mitochondrial genome of Dipodomys ordii that the first has the complete mitochondrial genome in the genus of Heteromyidae. The mitogenome is a circular molecule of 16 257 bp in length, containing 13 protein-coding genes, 2 ribosomal RNAs, 22 transfer RNAs and a putative displacement loop region. All protein-coding genes started with a traditional ATN codon and terminated with the mitochondria stop codon (TAA/TAG/AGA) or a single T base. The gene order and composition of mitogenome was similar to that of most other Sciurognathi species and its GC content was 36.73%. Thirteen protein-coding genes of D. ordii together with eight other closely species were used to construct the species phylogenetic tree for verification of the accuracy of new determined mitogenome sequences.

Rapid genetic restoration of a keystone species exhibiting delayed demographic response.

Genetic founder effects are often expected when animals colonize restored habitat in fragmented landscapes, but empirical data on genetic responses to restoration are limited. We examined the genetic response of banner-tailed kangaroo rats (Dipodomys spectabilis) to landscape-scale grassland restoration in the Chihuahuan Desert of New Mexico, USA. Dipodomys spectabilis is a grassland specialist and keystone species. At sites treated with herbicide to remove shrubs, colonization by D. spectabilis is slow and populations persist at low density for ≥10 years (≥6 generations). Persistence at low density and low gene flow may cause strong founder effects. We compared genetic structure of D. spectabilis populations between treated sites and remnant grasslands, and we examined how the genetic response to restoration depended on treatment age, area, and connectivity to source populations. Allelic richness and heterozygosity were similar between treated sites and remnant grasslands. Allelic richness at treated sites was greatest early in the restoration trajectory, and genetic divergence did not differ between recently colonized and established populations. These results indicated that founder effects during colonization of treated sites were weak or absent. Moreover, our results suggested founder effects were not mitigated by treatment area or connectivity. Dispersal is negatively density-dependent in D. spectabilis, and we hypothesize that high gene flow may occur early in the restoration trajectory when density is low. Our study shows genetic diversity can be recovered more rapidly than demographic components of populations after habitat restoration and that founder effects are not inevitable for animals colonizing restored habitat in fragmented landscapes.

Divergence in sink contributions to population persistence.

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.

Kangaroo rats revisited: re-evaluating a classic case of desert survival.

Kangaroo rats are the archetypical organisms for mammalian survival in North American deserts, yet there are contradictions in the data surrounding their physiology and ecology. The traditional view has been that these nocturnal rodents have little tolerance to high temperatures (e.g., >30°C), reside in cool, humid burrows to escape the heat of the day, and nearly exclusively rely on a dry, carbohydrate-rich diet from which they metabolically derive most of their water supply. To test this view, we measured the microclimates, activity, and diet of Merriam's kangaroo rats (Dipodomys merriami) from a xeric location within the center of the Sonoran Desert. We arrive at the following conclusions: 1. Burrows are much hotter during the summer than previously appreciated. For over 100 days of the year, soil temperatures exceed 30°C at depths to 2 m. For over 50 days, temperatures exceed 35°C at depths to 1.5 m. These high temperatures at such depths preclude kangaroo rats from locating to cool temperatures (e.g., <30°C) by burrowing. 2. Kangaroo rats remain in shallow burrows (<1 m) at relatively high ambient temperatures (>35°C) throughout the daytime during the summer instead of residing deep within the soil. This finding supports recent laboratory experiments that show kangaroo rats have much higher thermal tolerances than previously realized. 3. Kangaroo rats do not restrict their activity to the coolest periods of the night, but are active immediately following sundown, during the hottest time of the night. 4. Burrows are not persistently humid, but can be quite dry. 5. Insects and succulent vegetation constitute a significant portion of a kangaroo rat's diet and may be key to their survival in the hot desert environment.

Food caching and differential cache pilferage: a field study of coexistence of sympatric kangaroo rats and pocket mice.

Ecologists studying sympatric heteromyid rodents have sought evidence for species differences in primary foraging abilities and preferences and/or behavioural responses to predation risk in order to explain coexistence. The present field study was conducted to test the hypothesis that another factor may be involved, namely differences in caching patterns, which may result in differences in vulnerability to pilferage. We examined differences between kangaroo rats (Dipodomys merriami) and pocket mice (Chaetodipus spp.) in foraging, caching and pilferage behaviour. Specifically, we examined interactions at food patches, differential food caching patterns, and differential vulnerability to cache pilferage. Observations conducted at artificial seed patches showed that kangaroo rats dominated access to the patches by arriving and foraging first and by chasing pocket mice away. Individually provisioned pocket mice stored most seeds in underground burrows (larder hoarding), whereas kangaroo rats predominantly cached seeds in small, spatially dispersed caches in shallow pits in the surface of the sand (scatter hoarding). Pocket mice pilfered from each other as well as from the kangaroo rats, but the kangaroo rats rarely pilfered, and the only instance was from another kangaroo rat. Kangaroo rats and pocket mice were both vulnerable to cache pilferage. The results suggest that coexistence of kangaroo rats and pocket mice may be facilitated by a trade-off between primary harvest ability and the ability to exploit a resource that has been processed by another species, namely pilferage ability.

Mechanisms for the keystone status of kangaroo rats: graminivory rather than granivory?

Graminivory by kangaroo rats (Dipodomys spp.) was investigated as a potential mechanism for the keystone role of these rodents in the dynamics of desert grasslands. Experiments confirmed that Ord's kangaroo rats (Dipodomys ordii) cut and consumed a large proportion of the tillers of three Chihuahuan Desert tussock-forming grass species. Field observations indicated that the characteristically cut grass tillers were absent from all-rodent and medium-sized kangaroo rat exclosures, but were frequent in large-sized kangaroo rat and rabbit exclosures, indicating that the medium-sized kangaroo rats (D. ordii, D. merriami) were responsible for grass cutting. Tiller waste as a percentage of peak standing crop ranged from 7% in grassland habitats to 0.7% in Flourensia cernua shrubland. Of the 13 species of perennial, tussock-forming grasses measured, only one, Muhlenbergia porteri, had no tillers cut by kangaroo rats. This study demonstrates that the keystone role of kangaroo rats in Chihuahuan Desert grassland ecosystems is probably the result of their graminivory.