Tunnel-tube and Fourier methods for measuring three-dimensional medium reaction force in burrowing animals.

Subterranean digging behaviors provide opportunities for protection, access to prey, and predator avoidance for a diverse array of vertebrates, yet studies of the biomechanics of burrowing have been limited by the technical challenges of measuring kinetics and kinematics of animals moving within a medium. We describe a new system for measuring 3D reaction forces during burrowing, called a 'tunnel-tube', which is composed of two, separately instrumented plastic tubes: an 'entry tube' with no medium, in series with a 'digging tube' filled with medium. Mean reaction forces are measured for a digging bout and Fourier analysis is used to quantify the amplitude of oscillatory digging force as a function of frequency. In sample data from pocket gophers digging in artificial and natural media, the mean ground reaction force is constant, whereas Fourier analysis resolves a reduced amplitude of oscillatory force in the artificial medium with lower compaction strength.

Locomotory adaptations in entoptychine gophers (Rodentia: Geomyidae) and the mosaic evolution of fossoriality.

Pocket gophers (family Geomyidae) are the dominant burrowing rodents in North America today. Their fossil record is also incredibly rich; in particular, entoptychine gophers, a diverse extinct subfamily of the Geomyidae, are known from countless teeth and jaws from Oligocene and Miocene-aged deposits of the western United States and Mexico. Their postcranial remains, however, are much rarer and little studied. Yet, they offer the opportunity to investigate the locomotion of fossil gophers, shed light on the evolution of fossoriality, and enable ecomorphological comparisons with contemporaneous rodents. We present herein a quantitative study of the cranial and postcranial remains of eight different species of entoptychine gophers as well as many contemporary rodent species. We find a range of burrowing capabilities within Entoptychinae, including semifossorial scratch-digging animals and fossorial taxa with cranial adaptations to burrowing. Our results suggest the repeated evolution of chisel-tooth digging across genera. Comparisons between entoptychine gophers and contemporaneous rodent taxa show little ecomorphological overlap and suggest that the succession of burrowing rodent taxa on the landscape may have had more to do with habitat partitioning than competition.

Morphological adaptations for digging and climate-impacted soil properties define pocket gopher (Thomomys spp.) distributions.

Species ranges are mediated by physiology, environmental factors, and competition with other organisms. The allopatric distribution of five species of northern Californian pocket gophers (Thomomys spp.) is hypothesized to result from competitive exclusion. The five species in this environmentally heterogeneous region separate into two subgenera, Thomomys or Megascapheus, which have divergent digging styles. While all pocket gophers dig with their claws, the tooth-digging adaptations of subgenus Megascapheus allow access to harder soils and climate-protected depths. In a Northern Californian locality, replacement of subgenus Thomomys with subgenus Megascapheus occurred gradually during the Pleistocene-Holocene transition. Concurrent climate change over this transition suggests that environmental factors--in addition to soil--define pocket gopher distributional limits. Here we show 1) that all pocket gophers occupy the subset of less energetically costly soils and 2) that subgenera sort by percent soil clay, bulk density, and shrink-swell capacity (a mineralogical attribute). While clay and bulk density (without major perturbations) stay constant over decades to millennia, low precipitation and high temperatures can cause shrink-swell clays to crack and harden within days. The strong yet underappreciated interaction between soil and moisture on the distribution of vertebrates is rarely considered when projecting species responses to climatic change. Furthermore, increased precipitation alters the weathering processes that create shrink-swell minerals. Two projected outcomes of ongoing climate change--higher temperatures and precipitation--will dramatically impact hardness of soil with shrink-swell minerals. Current climate models do not include factors controlling soil hardness, despite its impact on all organisms that depend on a stable soil structure.

Thomas George Lee - Implantation and early development of North American rodents.

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.

Costs and benefits of pocket gopher foraging: linking behavior and physiology.

Animals can attain fitness benefits by maintaining a positive net energy balance, including costs of movement during resource acquisition and the profits from foraging. Subterranean rodent burrowing provides an excellent system in which to examine the effects of movement costs on foraging behavior because it is energetically expensive to excavate burrows. We used an individual-based modeling approach to study pocket gopher foraging and its relationship to digging cost, food abundance, and food distribution. We used a unique combination of an individual-based foraging-behavior model and an energetic model to assess survival, body mass dynamics, and burrow configurations. Our model revealed that even the extreme cost of digging is not as costly as it appears when compared to metabolic costs. Concentrating digging in the area where food was found, or area-restricted search (ARS), was the most energetically efficient digging strategy compared to a random strategy. Field data show that natural burrow configurations were more closely approximated by the animals we modeled using ARS compared to random diggers. By using behavior and simple physiological principles in our model, we were able to observe realistic body mass dynamics and recreate natural movement patterns.

Rodenticide grain bait ingredient acceptance by Norway rats (Rattus norvegicus), California ground squirrels (Spermophilus beecheyi) and pocket gophers (Thomomys bottae).

Vertebrate pest control in California is often accomplished through the use of rodenticide grain baits. These grain baits are composed of steam-rolled oats (SRO), a toxicant, an indicator dye and an oil combination. A series of tests were performed to determine the effects of various dye and oil formulations on acceptance of grain bait by Norway rats [Rattus norvegicus (Berk)], California ground squirrels [Spermophilus beecheyi (Richardson)] and pocket gophers (Thomomys bottae Eyd & Gerv). Seven different dyes, four oil formulations and clean (untreated) oats were tested for acceptance. The addition of the selected oils and dyes to grain resulted in no significant differences in consumption. This indicates that there is a wide variety of dyes that could be used in the formulation of rodenticides. These alternatives could aid in proper pesticide use, the deterrence of bait consumption by birds and possibly in ingredient adhesion to the finished bait.

Proximal cues of pocket gopher burrow plugging behavior: influence of light, burrow openings, and temperature.

Burrow plugging is readily observed among mammals adapted for digging (i.e., fossorial mammals) as they create and maintain their burrows. We investigated the influence of light, burrow openings, and thermal environment as cues of pocket gopher (Thomomys mazama, Thomomys talpoides) behavior. When given free access to light and no light during artificial-burrow preference trials, both Thomomys spp. consistently plugged (i.e., avoided) light treatments. Burrow openings did not notably affect plugging behavior of T. mazama. Gophers (T. talpoides) plugged the artificial burrows within the light and cold (7 degrees C) treatments, but not within the no-light, and 18 or 31 degrees C treatments when light and temperature were varied independently. Whereas the presence of light and low ambient temperatures induce burrow maintenance by pocket gophers, these cues help meliorate adverse conditions within subsurface environs.

Photoreceptors and photopigments in a subterranean rodent, the pocket gopher (Thomomys bottae).

Pocket gophers (Thomomys bottae) are rodents that spend much of their lives in near-lightless subterranean burrows. The visual adaptations associated with this extreme environment were investigated by making anatomical observations of retinal organization and by recording retinal responses to photic stimulation. The size of the eye is within the normal range for rodents, the lens transmits light well down into the ultraviolet, and the retina conforms to the normal mammalian plan. Electroretinogram recording revealed the presence of three types of photopigments, a rod pigment with a spectral peak of about 495 nm and two types of cone pigment with respective peak values of about 367 nm (UV) and 505 nm (medium-wavelength sensitive). Both in terms of responsivity to lights varying in temporal frequency and in response recovery following intense light adaptation, the cone responses of the pocket gopher are similar to those of other rodents. Labeling experiments indicate an abundance of cones that reach densities in excess of 30,000 mm-2. Cones containing UV opsin are found throughout the retina, but those containing medium-wavelength sensitive opsin are mostly restricted to the dorsal retina where coexpression of the two photopigments is apparently the rule. Rod densities are lower than those typical for nocturnal mammals.