Seasonal patterns of nitrogen utilization by pocket gophers, Thomomys bottae

Adult pocket gophers, fed diets of natural vegetation in the laboratory during summer and winter, remained in positive N balance in all feeding trials. Nitrogen budgets for free-ranging pocket gophers were calculated and revealed that total N influx exceeded 2400 mg N . kg-1 day-1 in summer and 2300 mg N . kg-1 day-1 in winter. Apparent N assimilation was 50 and 53% in summer and winter, respectively, and truly digestible N requirements were congruent to 680 mg N . kg-1 day-1. These budgets indicate that, in spite of seasonally-changing quality of available forage, pocket gophers remain in positive N balance, due largely to shifts in forage preference with season.     

Burrow structure and foraging costs in the fossorial rodent,Thomomys bottae

Summary A model for calculating the energy cost of burrowing by fossorial rodents is presented and used to examine the energetics of foraging by burrowing. The pocket gopher Thomomys bottae (Rodentia: Geomyidae) digs burrows for access to food. Feeding tunnels of Thomomys are broken into segments by laterals to the surface that are used to dispose of excavated soil. Energy cost of burrowing depends on both soil type and on burrow structure, defined by the length of burrow segments, angle of ascent of laterals, depth of feeding tunnels, and burrow diameter. In a desert scrub habitat, Thomomys adjust burrow segment length to minimize cost of burrowing. Observed segment lengths (mean=1.33 m) closely approximate the minimum-cost segment length of 1.22 m. Minimizing energy expended per meter of tunnel constructed maximizes efficiency of foraging by burrowing in the desert scrub. Burrow diameter and cost of burrowing increase with body size, while benefits do not, so foraging by burrowing becomes less enconomical as body size increases. Maximum possible body size of fossorial mammals depends on habitat productivity and energy cost of burrowing in local soils.     

Genome evolution in pocket gophers (genus Thomomys)

Heterochromatin is a dominant component of the genome in the bottae group of the pocket gopher genus Thomomys, having had a major role in the karyotypic evolution of member species. Heterochromatin characteristics of two subspecies of T. bottae and one of T. umbrinus were examined with fluorochrome dyes identifying presumptive GC- and AT-rich regions. In two karyotypic forms of T. b. fulvus and in T. umbrinus, chromatin that fluoresces brightly with chromomycin A₃ is also C-band positive, although not all heterochromatin fluoresces. However, in T. b. bottae, only euchromatic regions fluoresce brightly with chromomycin. Fluorescence patterns produced with DAPI are the reverse of the chromomycin banding in all karyotypic forms. Heterochromatin in these taxa is thus highly differentiated, exhibiting heterogeneity in staining characteristics, and presumably in underlying DNA sequences, both across the genome within a given chromosomal complement as well as among the different karyotypic races and species of the bottae group of pocket gophers.     

Genome evolution in pocket gophers (genus Thomomys)

Cellular DNA content (2 C-value) was measured by fluorescence flow cytometry of chromomycin-A₃ stained spleen cells in 2 subgenera, 5 species, and 21 subspecies of pocket gophers (genus Thomomys). The data indicate that, in Thomomys: (1) interspecific variation is extensive but, while some congeneric species differ by as much as 230%, others are identical in C-value; (2) intraspecific differentiation can be extensive with C-values differing by as much as 35%; and (3) populations of the same subspecies with apparently similar karyotypes can differ significantly in C-value. The implications of these results for hypotheses of the adaptive significance of C-value variation and genome evolution are discussed.     

Karyotypic variation following an elevational gradient in the pocket gopher,Thomomys bottae grahamensis Goldman

A system of non-Robertsonian chromosomal polymorphism is described for populations of the pocket gopher, Thomomys bottae grahamensis Goldman, from the Graham Mountains of southeastern Arizona. The polymorphism occurs in three of the four populations sampled, but the range of chromosomal variation within each population is less than for the whole sample. The diversity varies clinally with elevation and is strongly correlated with both habitat and moisture gradients. Each population is considered to be at an adaptive norm with respect to the occupied habitat, and the karyotypic changes are presumed to have played a role in this adaptability. Maintenance of the polymorphic conditions in each population is considered to be due to karyotypic stability (either frequency equilibrium or population monotypy) coupled with gene flow from adjacent populations.     

Geographic genetic architecture of pocket gopher (Thomomys bottae) populations in Baja California, Mexico

Phylogenetic analyses of complete mitochondrial cytochrome b sequences support the monophyly of pocket gopher (Thomomys bottae) populations from the 1000 km length of the Baja California peninsula of Mexico, relative to other geographical segments of the species range in western North America. The Baja California peninsula is an area that encompasses considerable ecomorphological and infraspecific diversity within this pocket gopher species. However, detailed population analyses encompassing 35 localities distributed over the southern half of the peninsula reveal only trivial phylogeographical structure. Rather, most of the 72 unique 500-base pair haplotypes examined from 142 individuals is restricted to single populations, although a few haplotypes are shared broadly across geography. Individual populations are typically comprised of haplotype sets from different branches in a network of relationships. Analysis of molecular variance (amova) indicates that approximately half of the total pool of variation is contained among individuals within local populations, and that only about 25% can be explained by the regional subdivisions of current subspecies distributions or physiographic realms. A hypothesized historical vicariant event that has been causally linked to the phylogeographical structure of other, codistributed species has had little influence on these pocket gopher populations, explaining only 13% of the total variation. The temporal depth, estimated by coalescence parameters, of the haplotype lineage in Baja California is relatively recent, approximately 300,000 generations; both the mismatch distribution of pairwise comparisons and a significantly positive exponential growth estimate support a recent history of expanding populations; but current, or recent past, migration estimates have remained small, are largely unidirectional from north to south, and weak isolation by distance is present. All data suggest that pocket gophers have relatively recently invaded the southern half of peninsular Baja California, with the genetic signature of expansion still evident but with sufficient time having lapsed to result in a weak isolation by distance pattern. The geographical assemblage of sampled populations thus appears as a meta-population, with limited gene flow contrasting with random haplotype loss due to drift in small, localized populations.     

Effects of the northern pocket gopher (Thomomys talpoides) on alpine soil characteristics, Niwot Ridge, CO

Effects of the northern pocket gopher (Thomomys talpoides) on surface soilcharacteristics were examined at the alpinesite of Niwot Ridge, CO. We measured erosionof soil from gopher mounds and compared thecharacteristics of gopher mound (disturbed) andundisturbed soils in two major plant communitytypes. Our measurements of erosion indicatelong-term susceptibility of gopher-disturbedsoils to redistribution by water and/or wind inthis ecosystem. Ecosystem heterogeneityintroduced by the gopher is reflected insignificantly lower SOM in gopher mounds thanin surrounding undisturbed soils, acharacteristic which appears to be causallyassociated with other effects of gopherdisturbance including changes in soil textureand significantly lower clays, total C, totalN, total P, and labile P. In contrast toplant-available P, NO₃ ^– was higherand steadily increased for the short term in both gopher mound soils and those beneath the mounds. These pools of NO₃ ^– thendecreased to pre-disturbance levels by thefollowing spring. Collectively our resultsindicate that, through the physicalmanipulation of soil and subsequent effects onsoil resources, the northern pocket gopherfunctions as an agent of increased ecosystemheterogeneity and soil mass and nutrientredistribution at Niwot Ridge.     

The relationship between genetic variability and growth rate among populations of the pocket gopher, Thomomys bottae

Perhaps the oldest unresolved debate inconservation genetics is whether geneticvariability matters – in other words, whetherrelatively low average genetic variationcontributes to deficits in individual andpopulation level vigor and fitness. Using astatistically powerful paired sampling designin which each of three pairs of populationsconsisted of one high genetic variability andone low genetic variability population from aparticular subspecies of the pocket gopher,Thomomys bottae, we tested the hypothesisthat individuals from populations with lowergenetic variability have lower growth rates (acommonly used surrogate for fitness) than thosefrom populations with higher variability. Wemeasured genetic variability using averageallozyme heterozygosity and two measures of DNAfingerprint band sharing (Jeffreys 33.15 andMS1 probes). The population rankings of thelevels of genetic variability among the threemeasures were concordant. The least squaresmean growth rate (controlling for sex,subspecies and initial mass) of gophers fromlow variability populations (0.41 ± 0.06g/day, n = 48) was less than half that ofgophers from high variability populations (1.04± 0.07 g/day, n = 45). This result lendscredence to the premise that differences inpopulation level genetic variability havesignificant fitness consequences andunderscores the importance of maintaininggenetic variability in managed populations.     

Phylogenetic structure of the Thomomys bottae–umbrinus complex in North America

The phylogeography of the Thomomys bottae–umbrinus complex in the United States and Mexico was assessed with sequences of the mitochondrial cytochrome b gene. These sequences were obtained from 225 individuals representing 108 locations over the range, including 56 sequences from GenBank. 110 (500 bp) sequences were used for Bayesian inference and neighbor-joining analyses, and 34 (1140 bp) specimens from the main clades obtained from the Bayesian inference were used in maximum-parsimony and maximum-likelihood analyses. The different analyses indicate significant variation within the species complex that averages 13% among major groups of genetic differences among Thomomys bottae–umbrinus. The overall pattern of geographic variation is not concordant with the current taxonomy. To the contrary, eight monophyletic groups are supported by all analyses and can be considered phylogenetic species. Overall divergence among these groups appears influenced by historical biogeographic events active during the Pliocene and Pleistocene.     

Electrophoretic characterization of the hemoglobins in pocket gophers of the genus Thomomys.

1. Hemoglobin phenotypes have been determined for closely related pocket gopher subspecies in the genus Thomomys from sea level and high altitude. 2. A single hemoglobin band was demonstrated for each gopher population with cellulose acetate membrane electrophoresis, with a mobility of 73% relative to human Hb-A. 3. Using polyacrylamide gel electrophoresis, a major Hb component (94% of the total Hb) showed a relative mobility of 68-69%; the minor Hb component showed a relative mobility of 74-75%. 4. The electrophoretic and physiological similarities of the gopher hemoglobins have persisted despite extreme differences in ambient PO2 in the habitats of each subspecies.     

Responses of pocket gophers (Thomomys bottae) to changes in diet quality

Summary We examined digestibility of dry matter, nutrients, and fiber, and food intake, metabolic fecal losses, weight change, and gut size of pocket gophers (Thomomys bottae) in relation to diet quality in the laboratory. Pocket gophers were maintained for 15–20 days on one of seven diets which contained from 18% to 56% neutral detergent fiber (NDF). NDF content of the diet was an excellent predictor of diet quality. Digestibility of dry matter, NDF, and nitrogen all decreased with increasing NDF content of the diet. In general, pocket gophers compensated for low diet quality by increasing dry matter intake, but those given high quality forage before the lowest quality diet reduced their intake. Thus, the response of pocket gophers to low quality diets may depend on their body condition. Because increased food intake resulted in increased total metabolic fecal losses and metabolic fecal nitrogen losses, decreasing food intake on low-quality diets may be advantageous. A further response of pocket gophers to decreased food quality was an increase in size of cecum and large intestine, suggesting that fermentation of cell walls became increasingly important as diet quality decreased.     

Phylogenetic structure among pocket gopher populations,genus Thomomys (Rodentia: Geomyidae), on the Baja California Peninsula

We determined the phylogenetic relationships, population history, and hierarchical structure of genetic variation in pocket gophers distributed on the Baja California Peninsula (BCP), based on extensive geographic sampling. Using a fragment of the mitochondrial gene cytochrome b (cyt b), we found three latitudinal structured geographic clades (northern, central, and southern). The northern clade occurs in the border area of the USA and the north of BCP, the central clade occurs from the peninsular highlands through the Central Desert of Baja California, and the southern clade is distributed south of the San Ignacio Lagoon. AMOVA showed that genetic variation is higher among clades (64%) than within populations (18.1%). The deepest divergence among clades is very shallow (～300 000 years), which suggests that climatic changes during the Pleistocene or some inhospitable habitats have affected the structure of this group, rather than influences from older marine transgressions. Phylogenetic groups disclosed by our results do not coincide with the current infraspecific classification; therefore, we propose a change of epithet for BCP gophers (Thomomys nigricans) and a new subspecific taxonomic arrangement with four subspecies: Thomomys nigricans anitae, Thomomys nigricans martirensis, Thomomys nigricans nigricans, and Thomomys nigricans russeolus. © 2013 The Linnean Society of London