Dipodomys Deserti Research Articles

Stability of kangaroo rat burrows in the Sonoran Desert: Evidence of biocementation

Kangaroo rats (Dipodomys deserti) construct complex burrow systems in loose desert sand that survive temperature and relative humidity fluctuations and storms. Animals that burrow in desert sand typically burrow in compacted sand, near plant roots, or when the soil is unsaturated. However, these processes are insufficient to explain tunnel stability of kangaroo rats. Our goal is to understand how kangaroo rat burrows remain stable in loose desert sand, intending to translate this knowledge to geotechnical engineering. A kangaroo rat habitat in the dunes of The Sonoran Desert, AZ, was selected for the study. Dynamic cone penetrometer tests performed at active, abandoned, and no-burrow sites demonstrated that the animals prefer loose sand for burrow construction. Soil samples collected from the burrows' ceilings, subsurface, and surface were characterized. Brazilian tensile strength test results showed that burrow soil has approximately 3 times greater tensile strength than the rest at dry state, which indicates increased interparticle attractive stress in burrow ceilings due to biocementation. Laboratory experiments, scanning electron microscopy, and confocal microscopy images showed that fungal and microbial biofilms provided 17 kPa increase in interparticle attractive stress at less than 1% biomass concentration, indicating potential to be used in soil improvement applications.

Scientists may have discovered why Speedy Gonzales is always fast

Almost every person has walked or run on the beach at some point in their life. Moving on loose sand feels more challenging than walking on a sidewalk. In general, walking on sand requires more work and energy and this is not unique to humans. Now, you may also be familiar with Speedy Gonzales, a Looney Tunes cartoon mouse who runs super-fast all the time. You may wonder how Speedy can always run so fast across all the different terrains that he encounters, but a very similar looking animal to Speedy Gonzales in real life, the kangaroo rat (Dipodomys deserti), may prove that the cartoon's versatility is not so far off reality.Some researchers have previously tested different types of animals, such as lizards and ostriches, running over solid surfaces versus sandy surfaces. This line of research motivated Joseph Hall (Washington State University, USA), Craig McGowan (University of Idaho, USA) and David Lin (Washington State University) to study a hopping animal, the kangaroo rat, to expand on our current understanding of the effects of different types of ground on animal movements. These little hopping animals are equally adept moving across rocky and sandy terrains, both of which exist in their natural desert habitat. So, the researchers cleverly developed the very first variable terrain rotatory treadmill and put these animals to the test hopping on a solid and a sandy surface while controlling the speed and recording video of the animals’ manoeuvres.However, before the kangaroo rats were allowed to hop in place, the team had to ensure that the level of sand on the treadmill was sufficient to prevent the animal from touching the solid base beneath. To do this they designed a device that can measure how far the rodents’ feet would penetrate the sand and determined that at least 1.5 cm of sand was sufficient.Although the six desert kangaroo rats had to move at five different speeds, ranging from 1.25 to 2.25 m s−1, Hall and colleagues only analysed the motion of the little rodents’ hind legs at 1.8 m s−1, the speed at which these animals tend to hop in the wild. Comparing the animals’ hopping styles on solid and sandy surfaces, they found no significant difference in the amount of time the animals’ feet were in contact with the ground or the distance covered during each hop. Also, the researchers found no meaningful difference in the rodent's speed and acceleration between the two different surfaces, although the animals were more crouched when hopping on sand.Even though the environment has induced changes in the running styles of some animals, it appears that kangaroo rats have evolved so that changes in the surface they hop around on does not affect their speed. They maintain very similar physical hindlimb movements regardless of the firmness of the surface that they are moving over, suggesting that they are finely tuned to their environment. And even though Speedy Gonzales is a runner and kangaroo rats hop, the discovery that kangaroo rats are perfectly tuned to move on hard and soft surfaces could hold the key to Speedy's super-fast reputation.

Comparison between the kinematics for kangaroo rat hopping on a solid versus sand surface.

In their natural habitats, animals move on a variety of substrates, ranging from solid surfaces to those that yield and flow (e.g. sand). These substrates impose different mechanical demands on the musculoskeletal system and may therefore elicit different locomotion patterns. The goal of this study is to compare bipedal hopping by desert kangaroo rats (Dipodomys deserti) on a solid versus granular substrate under speed-controlled conditions. To accomplish this goal, we developed a rotary treadmill, which is able to have different substrates or uneven surfaces. We video recorded six kangaroo rats hopping on a solid surface versus sand at the same speed (1.8 m s−1) and quantified the differences in the hopping kinematics between the two substrates. We found no significant differences in the hop period, hop length or duty cycle, showing that the gross kinematics on the two substrates were similar. This similarity was surprising given that sand is a substrate that absorbs mechanical energy. Measurements of the penetration resistance of the sand showed that the combination of the sand properties, toe-print area and kangaroo rat weight was probably the reason for the similarity.

Elastic energy storage across speeds during steady-state hopping of desert kangaroo rats (Dipodomys deserti).

Small bipedal hoppers, including kangaroo rats, are not thought to benefit from substantial elastic energy storage and return during hopping. However, recent species-specific material properties research suggests that, despite relative thickness, the ankle extensor tendons of these small hoppers are considerably more compliant than had been assumed. With faster locomotor speeds demanding higher forces, a lower tendon stiffness suggests greater tendon deformation and thus a greater potential for elastic energy storage and return with increasing speed. Using the elastic modulus values specific to kangaroo rat tendons, we sought to determine how much elastic energy is stored and returned during hopping across a range of speeds. In vivo techniques were used to record tendon force in the ankle extensors during steady-speed hopping. Our data support the hypothesis that the ankle extensor tendons of kangaroo rats store and return elastic energy in relation to hopping speed, storing more at faster speeds. Despite storing comparatively less elastic energy than larger hoppers, this relationship between speed and energy storage offers novel evidence of a functionally similar energy storage mechanism, operating irrespective of body size or tendon thickness, across the distal muscle-tendon units of both small and large bipedal hoppers.

Plantar flexor muscles of kangaroo rats (Dipodomys deserti) shorten at a velocity to produce optimal power during jumping.

The musculotendon work contributions across all joints during jumping by kangaroo rats are not well understood. Namely, measures of external joint work do not provide information on the contributions from individual muscles or in-series elastic structures. In this study, we examined the functional roles of a major ankle extensor muscle, the lateral gastrocnemius (LG), and a major knee extensor muscle, the vastus lateralis (VL), through in vivo sonomicrometry and electromyography techniques, during vertical jumping by kangaroo rats. Our data showed that both muscles increased shortening and activity with higher jumps. We found that knee angular velocity and VL muscle shortening velocity were coupled in time. In contrast, the ankle angular velocity and LG muscle shortening velocity were decoupled, and rapid joint extension near the end of the jump produced high power outputs at the ankle joint. Further, the decoupling of muscle and joint kinematics allowed the LG muscle to prolong the period of shortening velocity near optimal velocity, which likely enabled the muscle to sustain maximal power generation. These observations were consistent with an LG tendon that is much more compliant than that of the VL.

How to Stick the Landing: Kangaroo Rats Use Their Tails to Reorient during Evasive Jumps Away from Predators.

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.

Elemental and radionuclide exposures and uptakes by small rodents, invertebrates, and vegetation at active and post-production uranium mines in the Grand Canyon watershed

The effects of breccia pipe uranium mining in the Grand Canyon watershed (Arizona) on ecological and cultural resources are largely unknown. We characterized the exposure of biota to uranium and co-occurring ore body elements during active ore production and at a site where ore production had recently concluded. Our results indicate that biota have taken up uranium and other elements (e.g., arsenic, cadmium, copper, molybdenum, uranium) from exposure to ore and surficial contamination, like blowing dust. Results indicate the potential for prolonged exposure to elements and radionuclides upon conclusion of active ore production. Mean radium-226 in deer mice was up to 4 times greater than uranium-234 and uranium-238 in those same samples; this may indicate a potential for, but does not necessarily imply, radium-226 toxicity. Soil screening benchmarks for uranium and molybdenum and other toxicity thresholds for arsenic, copper, selenium, uranium (e.g., growth effects) were exceeded in vegetation, invertebrates, and rodents (Peromyscus spp., Thomomys bottae, Tamias dorsalis, Dipodomys deserti). However, the prevalence and severity of microscopic lesions in rodent tissues (as direct evidence of biological effects of uptake and exposure) could not be definitively linked to mining. Our data indicate that land managers might consider factors like species, seasonal changes in environmental concentrations, and bioavailability, when determining mine permitting and remediation in the Grand Canyon watershed. Ultimately, our results will be useful for site-specific ecological risk analysis and can support future decisions regarding the mineral extraction withdrawal in the Grand Canyon watershed and elsewhere.

Caching Propensities and Effectiveness of Five Coexisting Heteromyid Rodent Species as Dispersers of Indian Ricegrass (Achnatherum hymenoides) Seeds

Recruitment of Indian ricegrass (Achnatherum hymenoides) seedlings occurs primarily from surface seed caches made by scatter-hoarding desert rodents of the family Heteromyidae. We used radiolabeled Indian ricegrass seeds in field seed-caching experiments with 5 coexisting heteromyid species (Dipodomys deserti, D. merriami, D. microps, Microdipodops pallidus, and Perognathus longimembris) to compare their potential effectiveness as dispersers. Dipodomys microps individuals deposited seeds almost exclusively in larders rather than in scatter hoards, implying that this species is unlikely to be an important seed disperser. Among the other species, individual caches made by larger species had more seeds than those made by smaller species, but smaller species made more caches. At the level of individual animals, M. pallidus was the most effective disperser in a quantitative sense; they made more caches than other species tested and placed fewer excess seeds in caches relative to optimal cache sizes for Indian ricegrass seedling establishment. However, because D. merriami individuals were considerably more abundant at the study site than other species and were also avid scatter hoarders, D. merriami is likely to be the most quantitatively effective disperser of Indian ricegrass seeds at the species level. Ranking species according to qualitative effectiveness (e.g., by considering effects such as the caching microsite on seedling establishment) was more ambiguous. For example, P. longimembris made relatively shallow caches that most closely match optimal planting depths for Indian ricegrass seedling emergence, but such shallow caches are probably more likely than deeper caches to be discovered by foraging rodents before they can germinate. With the possible exception of D. microps, any of the species we tested may be effective dispersers of Indian ricegrass seeds.

Derived muscle arrangements and their shared innervation patterns of external and internal cheek pouches in rodents.

Cheek pouches have evolved from the oral cavity in rodents and act as temporary food storage repositories. There are two types of opening, internal and external. Details about the complex cutaneous muscles controlling the pouches have still not been fully elucidated. To understand the shared and derived traits of the muscles surrounding the cheek pouch and their innervation, we carried out an evolutionary morphological study using two desert kangaroo rats (Dipodomys deserti) and three plains pocket gophers (Geomys bursarius) from each of the two families equipped with external cheek pouches, and four Syrian hamsters (Mesocricetus auratus) with internal cheek pouches. The most conspicuous derived trait of the muscles between the external and internal cheek pouches, the sphincter sacculi, surrounds almost all of the edge of the outer entrance of the pouch. It is present in both species with external pouches, but not in hamsters, which have internal pouches. Our neurological findings demonstrate that most pouch muscles are innervated by both the facial and the cervical nerves, regardless of the pouch type. In these dually innervated muscles, the ventromedial part of the muscles tends to be innervated dominantly or uniquely by the cervical nerves, which usually enter from the superficial or lateral aspect. As a trait shared with the cervical nerves innervating the propatagial muscles in aerodynamic mammals such as bats and flying squirrels, and panniculus carnosus in most mammals, our neurological evidence suggests that the cervical nerve has the potential to innervate derived cutaneous muscles in the cervicofacial region.

Managing predators: The influence of kangaroo rat antipredator displays on sidewinder rattlesnake hunting behavior

AbstractUpon sensing predators in their vicinity, many prey species perform antipredator displays that are thought to provide information to the predator that deters it from attacking (predator‐deterrent signals). These displays can be complex, incorporating a variety of signaling elements as well as direct physical harassment of the predator. Although the display behaviors in these communication systems are often well characterized, evidence of the efficacy of these displays in deterring predators is limited due to the challenges associated with studying free‐ranging predators. Here, we examine how the anti‐snake signals of the desert kangaroo rat (Dipodomys deserti) influence the ambush hunting behaviors of sidewinder rattlesnakes (Crotalus cerastes). We found that, although desert kangaroo rats incorporate a number of signal elements into their antipredator display, only sand kicking behavior was a significant factor in motivating sidewinder rattlesnakes to cease hunting: high rates of sand kicking led to early abandonment of ambush coils. These results indicate that anti‐snake displays of small mammals may be especially effective at mitigating the threat posed by rattlesnakes when those displays incorporate physical harassment as well as signaling.

Determinants of predation success: How to survive an attack from a rattlesnake

Abstract The selection pressures that arise from capturing prey and avoiding predators are some of the strongest biotic forces shaping animal form and function. Examining how performance (i.e., athletic ability) affects the outcomes of encounters between free‐ranging predators and prey is essential for understanding the determinants of predation success rates and broad scale predator–prey dynamics, but quantifying these encounters in natural situations is logistically challenging. The goal of our study was to examine how various metrics of predator/prey performance determine predation success by studying natural predator–prey interactions in the field with minimal manipulation of the study subjects. We used high‐speed video recordings of free‐ranging sidewinder rattlesnakes (predator, Crotalus cerastes) and desert kangaroo rats (prey, Dipodomys deserti) to study how performance at various stages of their encounters alters the outcome of their interactions. We found that predation success depends on (a) whether the rattlesnake struck accurately, (b) if the rattlesnake strike was accurate, the reaction time and escape manoeuvres of the kangaroo rat, and (c) if the kangaroo rat was bitten, the ability of the kangaroo rat to use defensive manoeuvres to avoid subjugation by the snake. The results of our study suggest that the role of performance in predator–prey interactions is complex, and the determinants of predation success are altered by both predator and prey at multiple stages of an interaction in ways that may not be apparent in many experimental contexts. A plain language summary is available for this article.

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.

Functional capacity of kangaroo rat hindlimbs: adaptations for locomotor performance.

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. Lfibre) 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.

Recent interactions with snakes enhance escape performance of desert kangaroo rats (Rodentia: Heteromyidae) during simulated attacks

Recent interactions with snakes enhance escape performance of desert kangaroo rats (Rodentia: Heteromyidae) during simulated attacks

Avoiding the serpent's tooth: predator–prey interactions between free-ranging sidewinder rattlesnakes and desert kangaroo rats

Many species perform complex antipredator displays that deter attacks by informing predators that continued attempts at prey capture will be costly. However, because of the difficulties in studying the behaviour of free-ranging predators, we have a limited understanding of how predators respond to those signals. Here, we took advantage of our ability to quantify predatory behaviours of free-ranging sidewinder rattlesnakes, Crotalus cerastes , to examine the influence of anti-snake behaviours performed by desert kangaroo rats, Dipodomys deserti . We recorded natural encounters and quantified the predator-deterrent behaviours displayed by the kangaroo rats, as well as any strikes performed by the rattlesnakes and whether the strikes were successful. We found that predator-deterrent signalling significantly reduced the probability that a rattlesnake would strike. This was most likely due to the ability of kangaroo rats to mobilize extremely rapid evasive leaps; even rats that appeared unaware of the snakes were almost always able to avoid rattlesnake strikes. The degree of effectiveness of this evasive leaping in countering rattlesnake predation was unexpected and indicates that this may be a rich system for exploring the biomechanics of extreme physical performance in a naturalistic context.

Comparison of anti-snake displays in the sympatric desert rodentsXerospermophilus tereticaudus(round-tailed ground squirrels) andDipodomys deserti(desert kangaroo rats)

Several lineages of small mammals frequently preyed upon by snakes have evolved snake-specific signals and displays that they use in an attempt to deter predation. Although detailed studies have been conducted on the form and function of these behaviors in a few key species, limited work has been done that directly compares behaviors between species, and none that we know of between species of separate lineages. Representative species displaying these behaviors exhibit broad overlap in geographic range, and they often occur in sympatry with each other, where they are preyed upon by the same local guild of snakes. Thus, there is potential for these independently evolved signals to exhibit some degree of convergence in sympatry, reinforcing each other in a fashion similar to what has been documented for aposematic mimicry complexes. Here, we provide the 1st direct quantitative comparison of anti-snake signaling behavior in sympatric small mammals from separate lineages: Dipodomys deserti (Heteromyidae) and Xerospermophilus tereticaudus (Sciuridae). Although some aspects of the displays of squirrels and kangaroo rats were similar (e.g., use of foot drumming, time spent investigating snake), others were not (e.g., willingness to approach closely, tail flagging, sand kicking, jump back displays). Although further research will be necessary to test directly the possibility of convergence, the forms of the signals are disparate enough that it is unlikely that the displays represent an example of convergent mimicry. Our results highlight the potential for further comparative work regarding the evolution of form and function of anti-snake signaling in small mammals.

Genetic consequences of postglacial range expansion in two codistributed rodents (genus Dipodomys) depend on ecology and genetic locus.

How does range expansion affect genetic diversity in species with different ecologies, and do different types of genetic markers lead to different conclusions? We addressed these questions by assessing the genetic consequences of postglacial range expansion using mitochondrial DNA (mtDNA) and nuclear restriction site-associated DNA (RAD) sequencing in two congeneric and codistributed rodents with different ecological characteristics: the desert kangaroo rat (Dipodomys deserti), a sand specialist, and the Merriam's kangaroo rat (Dipodomys merriami), a substrate generalist. For each species, we compared genetic variation between populations that retained stable distributions throughout glacial periods and those inferred to have expanded since the last glacial maximum. Our results suggest that expanded populations of both species experienced a loss of private mtDNA haplotypes and differentiation among populations, as well as a loss of nuclear single-nucleotide polymorphism (SNP) private alleles and polymorphic loci. However, only D.deserti experienced a loss of nucleotide diversity (both mtDNA and nuclear) and nuclear heterozygosity. For all indices of diversity and differentiation that showed reduced values in the expanded areas, D.deserti populations experienced a greater degree of loss than did D.merriami populations. Additionally, patterns of loss in genetic diversity in expanded populations were substantially less extreme (by two orders of magnitude in some cases) for nuclear SNPs in both species compared to that observed for mitochondrial data. Our results demonstrate that ecological characteristics may play a role in determining genetic variation associated with range expansions, yet mtDNA diversity loss is not necessarily accompanied by a matched magnitude of loss in nuclear diversity.

P.1.h.024 3,4-Methylenedioxyethylamphetamine (MDEA) provokes changes in the sequential structure of the agonistic behaviour of mice

Kangaroo rats develop spongiform degeneration of the central auditory system similar to that seen in the gerbil. Light microscopic and transmission electron microscopic study of the cochlear nucleus and auditory nerve root (ANR) of Dipodomys deserti and D. merriami show that spongiform lesions develop in dendrites and oligodendrocytes of the cochlear nucleus and in oligodendrocytes of the ANR that are morphologically indistinguishable from those extensively described in the Mongolian gerbil, Meriones unguiculatus. As in Mongolian gerbils, the spongiform degeneration in Dipodomys were much more numerous in animals continually exposed to modest levels of low-frequency noise ( < 75 dB SPL). The kangaroo rats with extensive spongiform degeneration also show slightly, but significantly, elevated auditory brainstem evoked response (ABR) thresholds to low-frequency stimuli, a result also found in Mongolian gerbils. These results suggest that the elevated ABR thresholds may be the result of spongiform degeneration. Because low-frequency noise-induced spongiform degeneration has now been shown in the cochlear nucleus of animals from separate families of Rodentia (Heteromyidae and Muridae), the possibility should be investigated that similar noise-induced degenerative changes occur in the central auditory system of other mammals with good low-frequency hearing.

Collision-based mechanics of bipedal hopping

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.

Kinematic analysis of an appetitive food-handling behavior: the functional morphology of Syrian hamster cheek pouches

Prodigious food hoarding in Syrian hamsters Mesocricetus auratus Waterhouse is strongly linked to appetite and is made possible by large internal cheek pouches. We provide a functional analysis of the cheek pouch and its associated retractor muscle. Frame-by-frame analysis of videotaped pouch-filling behavior revealed multiple jaw cycles for each food item pouched and the use of more jaw cycles to pouch large food items ( approximately 2.5 g chow pellets) than small (corn kernels or sunflower seed with husks). These results stand in contrast to previously reported pouching kinematics in the externally pouched Dipodomys deserti, which uses only one jaw cycle per pouching event. Comparison of pouching and mastication in the same individuals also suggests that in Syrian hamsters, feeding jaw cycles are modulated to accommodate pouch filling primarily by the addition of a pause between fast open and fast close phases, which we call ;gape phase'. Contrary to previous assertions, the retractor muscle does not merely provide structural support for the full pouch during locomotion. Video analysis of ten hamsters with unilaterally denervated retractor muscles and electrophysiological study of an anaesthetized subject confirmed that retractor muscle activity during pouch filling increases pouching efficiency for food items subsequent to the first.