Temporal and spatial dynamics of competitive parapatry in chewing lice

We synthesize observations from 1979 to 2016 of a contact zone involving two subspecies of pocket gophers (Thomomys bottae connectens and T. b. opulentus) and their respective chewing lice (Geomydoecus aurei and G. centralis) along the Río Grande Valley in New Mexico, U.S.A., to test predictions about the dynamics of the zone. Historically, the natural flood cycle of the Rio Grande prevented contact between the two subspecies of pocket gophers. Flood control measures completed in the 1930s permitted contact, thus establishing the hybrid zone between the pocket gophers and the contact zone between their lice (without hybridization). Since that time, the pocket gopher hybrid zone has stabilized, whereas the northern chewing louse species has replaced the southern louse species at a consistent rate of ~150 m/year. The 0.2–0.8 width of the replacement zone has remained constant, reflecting the constant rate of chewing louse species turnover on a single gopher and within a local pocket gopher population. In contrast, the full width of the replacement zone (northernmost G. centralis to southernmost G. aurei) has increased annually. By employing a variety of metrics of the species replacement zone, we are better able to understand the dynamics of interactions between and among the chewing lice and their pocket gopher hosts. This research provides an opportunity to observe active species replacement and resulting distributional shifts in a parasitic organism in its natural setting.     

Simulation models of the interactions between herbivore foraging strategies, social behavior, and plant community dynamics

Herbivory often operates through a feedback in which herbivores affect the success and location of plants, which in turn affects the foraging behavior of animals. Factors other than food, such as social behavior, may influence the interactions between herbivores and the plants they consume. We used a simulation model to compare the effects of foraging and social behavior on plant distribution and foraging efficiency by gophers (Thomomys bottae) in a system characteristic of California grasslands. In this system, annual forbs are the preferred food items, and their abundance increases in areas disturbed by gopher burrowing. In addition, gopher social interactions generate buffer zones between adjacent burrows. During the first year of the simulations, before gophers affected the plant community, feeding efficiency declined with increased gopher density. However, after 40 yr, annual plant abundance increased with increasing gopher density, yielding higher maximum gopher density and per capita foraging efficiency. Conversely, increased width of the buffer zones lowered maximum gopher density and annual plant abundance resulting in lower feeding efficiency. In addition, the compact burrow structure of gophers employing an area-restricted search strategy allowed a higher density of gophers to coexist, resulting in higher annual plant abundance and higher per capita food-capture rates.     

Development and evolution of the carotid circulation in geomyoid rodents in relationship to their craniomorphology

The diversity in cranial morphology of living geomyoids, including pocket gophers (Thomomys), spiny pocket mice (Heteromys and Liomys), desert pocket mice (Chaetodipus and Perognathus), and kangaroo rats and mice (Dipodomys and Microdipodops) is accompanied by only a few differences in their cephalic arterial circulation. The principal difference is the origin of the pterygopalatine artery, which serves the orbit and rostrum. In dipodomyines and perognathines it originates as a stapedial branch of the internal carotid artery and passes through the middle ear en route to the braincase. In geomyines and heteromyines it originates as an internal maxillary branch of the external carotid artery and enters the braincase directly. Either geomyines and heteromyines are convergent in this respect, or the currently recognized family Heteromyidae is paraphyletic. The stapedial artery is the same size in early embryos of T. bottae and D. merriami, but in the former species it fails to grow and disappears in juveniles. Comparison of developmental series of D. merriami and T. bottae revealed that the decline of the artery in the latter species is preceded by a greater degree of arterial coarctation, or narrowing, as it passes though the developing stapes. The loss of the stapedial artery is correlated with an enlarged masseter profundus muscle in T. bottae and with an unusually small stapes in T. bottae, H. desmarestianus, and L. salvini. I hypothesize that the primitive condition for geomyoids is the presence of both stapedial and internal maxillary arteries, that the stapedial artery was lost in geomyines and heteromyines because of the constraint on its size posed by the stapes, and that the stapedial artery was retained in dipodomyines because enlargement of the stapes accompanied bullar inflation in these taxa and lifted the constraint on the size of the stapedial artery.     

Evolutionary and ecological response of pocket gophers (Thomomys talpoides) to late-Holocene climatic change

Late-Holocene evolutionary and ecological response of pocket gophers ( Thomomys talpoides ) and other species to climatic change is documented by mammalian fossils from Lamar Cave, a palaeontological site in northern Yellowstone National Park. Pocket gophers illustrate ecological sensitivity to a series of mesic to xeric climatic excursions in the sagebrush-grassland ecotone during the last 3200 years, increasing in abundance during mesic intervals, and declining in abundance during xeric intervals. Four other small mammal taxa (Microtus sp., Peromyscus maniculatus, Neotoma cinerea and Spermophilus armatus) also show response to climatic change, increasing or decreasing in abundance in accordance with their preferred habitat requirements. To determine evolutionary response to climate, two craniodental characters for the northern pocket gopher ( Thomomys talpoides ) are investigated in modern representatives within a 400 km radius of Lamar Cave and then tracked through the time spanned by the fossils. One morphologic character shows that variation in body size, primarily a plastic response to the environment, demonstrates few taxonomically consistent patterns of geographic variation across 39 modern localities. In contrast, the other character indicates genetic relatedness within subspecies. Stasis in the genetically controlled character indicates that the same subspecies of pocket gopher ( T. talpoides tenellus ) has occupied northern Yellowstone for at least 3200 years. However, T. t. tenellus does show plastic response to climatic change because pocket gophers during the Medieval Warm Period were smaller than at any other time spanned by the deposit.     

Photoreceptors and photopigments in a subterranean rodent,the pocket gopher (<Emphasis Type="Italic">Thomomys bottae</Emphasis>)

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.     

Fluctuating asymmetry and allozymic heterozygosity among natural populations of pocket gophers (Thomomys bottae)

Fluctuating asymmetry, or random deviations from bilateral symmetry, has been widely used as a measure of developmental stability. The relationship between fluctuating asymmetry (FA) and allozymic heterozygosity was evaluated using 18 natural populations of pocket gophers ( Thomomys bottae ). Heterozygosity in local populations of pocket gophers ranges over more than an order of magnitude (1.5—18.4%), making this burrowing rodent particularly apt for such studies. Two measures of FA in mensural skull characters were examined: absolute deviations between left and right sides and the variance of signed differences. After log transformations, levels of FA among individuals and populations were not related to size. Repeated-measures analyses of variance showed that FA was significant relative to measurement error, both across populations and within them. Asymmetries of different characters were uncorrelated, despite positive significant correlations among the characters themselves. FA levels varied only slightly among populations of gophers, and this variation was not significant for most characters. FA levels of populations were not correlated with allozymic heterozygosity, and analyses of variance in FA employing heterozygosity were not significant. Heterozygosity levels in these rodents appear more strongly related to aspects of population history (especially effective size and gene flow) than to developmental stability. Because so many genomic and environmental factors can affect morphological variation, caution is needed in interpreting correlations between genetic and phenetic variation.     

Phylogenetic evidence of mitochondrial DNA introgression among pocket gophers in New Mexico (family Geomyidae)

Mitochondrial DNA (mtDNA) variation in the cytochrome b gene was determined for two divergent taxa of pocket gophers, Thomomys bottae actuosus and T. b. ruidosae . These two taxa hybridize in a narrow contact zone, but introgression of nuclear markers such as allozymes or chromosomes does not extend much beyond the hybrid zone (Patton et al. 1979). We found that despite their distinctness, the two subspecies shared very similar mtDNA haplotypes. By a comparison of phylogenetic histories derived from nuclear markers (allozymes) and from mtDNA haplotypes sampled in different populations of T. bottae from New Mexico, we show that apparent similarity is due to an introgression of T. b. ruidosae mtDNA into T. b. actuosus nuclear background. Evidence of introgression is not limited to the present-day contact zone between these two taxa, but extends at least 75 km away from it. The actuosus haplotype coexists along with the ruidosae mtDNA in the Gallinas Mts., which are inhabited by otherwise pure T. b. actuosus , while further north only typical actuosus haplotypes were detected. Of several potential mechanisms which could lead to such a geographical pattern of variation, we argue that a combination of range shifts due to climatic fluctuations, and genetic drift are most likely. Horizontal gene transfers due to hybridization are historical events which seem rather common among pocket gophers. Although they can be identified with careful phylogenetic study using independent data sets, the potential for misinterpreting a gene tree as an organismal tree is great in this and other groups of animals.     

Two new species ofLitomosoides (Nematoda: Onchocercidae) from pocket gophers (Rodentia: Geomyidae) in Colorado

Summary Two new species of nematodes representing the genusLitomosoides were obtained from pocket gophers collected in the early 1960s and during 1982 in Colorado. Individuals ofLitomosoides thomomydis n. sp. were recovered from the abdominal cavities of two species of pocket gopher,Thomomys talpoides (Richardson) andT. bottae (Eydoux & Gervais). Nematodes representingL. westi n. sp. were recovered from the abdominal and pleural cavities of the plains pocket gopherGeomys bursarius (Shaw) from eastern Colorado. These two species of filarioid nematodes appear to be restricted in geographical range to those pocket gophers inhabiting the central grasslands and Rocky Mountain region of the United States and Canada. ac]19851217     

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.     

Oxidative activity and Delta(Psi) of liver mitochondria of active and hibernating ground squirrels during various incubation conditions]

A comparative analysis of delta psi of liver mitochondria of active and hibernating gophers was carried out, and the effect of the decrease of the medium tonicity on the oxidase activity and delta psi of liver mitochondria of both animal groups was studied. The delta psi of hibernating gopher liver mitochondria was shown to be lower than that of active animals mitochondria and displayed a higher sensitivity to the uncoupler and some different restoration dynamics after the decline following the ADP addition. Swelling of liver mitochondria of hibernating gophers in hypotonic media or in media containing potassium acetate and a 3-fold freeze-thawing procedure resulted in the enhancement of the oxidase activities accompanied (in case of hypotonicity) by an delta psi increase and a decrease of its sensitivity to the uncoupler, 2.4-dinitrophenol. Inhibition of phospholipase A2 prevented the enhancement of the oxidase activity in all cases and the delta psi increase in hypotonic media.     

Cophylogeny on a fine scale: Geomydoecus chewing lice and their pocket gopher hosts, Pappogeomys bulleri

Many species of pocket gophers and their ectoparasitic chewing lice have broadly congruent phylogenies, indicating a history of frequent codivergence. For a variety of reasons, phylogenies of codiverging hosts and parasites are expected to be less congruent for more recently diverged taxa. This study is the first of its scale in the pocket gopher and chewing louse system, with its focus entirely on comparisons among populations within a single species of host and 3 chewing louse species in the Geomydoecus bulleri species complex. We examined mitochondrial DNA from a total of 46 specimens of Geomydoecus lice collected from 11 populations of the pocket gopher host, Pappogeomys bulleri. We also examined nuclear DNA from a subset of these chewing lice. Louse phylogenies were compared with a published pocket gopher phylogeny. Contrary to expectations, we observed a statistically significant degree of parallel cladogenesis in these closely related hosts and their parasites. We also observed a higher rate of evolution in chewing louse lineages than in their corresponding pocket gopher hosts. In addition, we found that 1 louse species (Geomydoecus burti) may not be a valid species, that subspecies within G. bulleri are not reciprocally monophyletic, and that morphological and genetic evidence support recognition of a new species of louse, Geomydoecus pricei.     

AGE AND MOVEMENT OF A HYBRID ZONE: IMPLICATIONS FOR DISPERSAL DISTANCE IN POCKET GOPHERS AND THEIR CHEWING LICE

Historical flood records for the Rio Grande Valley of New Mexico suggest that a pocket gopher (Thomomys bottae) hybrid zone previously thought to be 10,000 years old may actually be closer to 50 years old. Measured zone width (defined genetically) is consistent with the hypothesis of recent contact, if we assume a reasonable dispersal distance of approximately 400 m/year for pocket gophers. A five-year study of movement of the contact zone between the two species of chewing lice that parasitize these pocket gophers also is consistent with the hypothesis of recent origin of the zone.     

Area-restricted search by the plains pocket gopher (Geomys bursarius) in tallgrass prairie habitat

Because pocket gophers have the high energetic cost of excavatingburrows and an inability to detect distant food items throughthe soil, I hypothesized that individuals within establishedburrow systems would use area-restricted search as a foragingstrategy. To examine this hypothesis I compared gopher foragingeffort over a 10-month period between areas in which overallplant densities were experimentally varied. Gophers expendedapproximately 50% of their foraging effort in areas with thehighest plant density, even though these made up only 33% ofthe available area in experimental plots. In large, griddedareas sampled for an entire season as well as in small areasin which gophers foraged for less than 1 week, gopher foragingeffort was related to the density of a single leguminous plantspecies, Psoralea argophylla. In small plots where this plantspecies was at high density, gophers created more tunnel branches,thereby intensifying their search effort. Thus, area-restrictedsearch appears to increase the rate of encounter with the patchilydistributed Psoralea plants.     

Molecular evidence for mating asymmetry and female choice in a pocket gopher (Thomomys) hybrid zone

This paper presents circumstantial evidence that the mating system of the North American pocket gophers (Rodentia: Geomyidae) is a promiscuous one, with female choice at its base. A molecular marker (a length variant in the mitochondrial Control region [D-loop]) is used to show mating asymmetry in a hybrid zone between the species Thomomys bottae and Thomomys townsendii in north-eastern California. All hybrids result from a bottae mother × townsendii father cross. Because of significant differences in body size and resulting burrow diameter, bottae females must have actively sought their respective townsendii mates for the asymmetry in mating to have occurred, signalling female choice in these subterranean mammals that are otherwise characterized by exclusive-use territories, skewed adult sex ratio in favour of females, and high variance in male reproductive success.     

Impact of pocket gopher disturbance on plant species diversity in a shortgrass prairie community

Summary We examined the impact of pocket gopher disturbances on the dynamics of a shortgrass prairie community. Through their burrowing activity, pocket gophers (Thomomys bottae) cast up mounds of soil which both kill existing vegetation and create sites for colonization by competitively-inferior plant species. Three major patterns emerge from these disturbances: First, we show that 10 of the most common herbaceous perennial dicots benefit from pocket gopher disturbance; that is, a greater proportion of seedlings are found in the open space created by pocket gopher disturbance than would be expected based on the availability of disturbed habitat. Additionally, these seedlings exhibited higher growth rates than adjacent seedlings of the same species growing in undisturbed habitat. Second, we tested two predictions of the Intermediate Disturbance Hypothesis and found that species diversity was greatest for plots characterized by disturbances of intermediate age. However, we did not detect significant differences in diversity between plots characterized by intermediate and high levels of disturbance, indicating that many species are adapted to or at least tolerant of high levels of disturbance. Third, we noted that the abundance of grasses decreased with increasing disturbance, while the abundance of dicots increased with increasing disturbance.     

Host behaviour drives parasite genetics at multiple geographic scales: population genetics of the chewing louse,Thomomydoecus minor

Pocket gophers and their symbiotic chewing lice form a host–parasite assemblage known for a high degree of cophylogeny, thought to be driven by life history parameters of both host and parasite that make host switching difficult. However, little work to date has focused on determining whether these life histories actually impact louse populations at the very fine scale of louse infrapopulations (individuals on a single host) at the same or at nearby host localities. We used microsatellite and mtDNA sequence data to make comparisons of chewing‐louse (Thomomydoecus minor) population subdivision over time and over geographic space where there are different potential amounts of host interaction surrounding a zone of contact between two hybridizing pocket‐gopher subspecies. We found that chewing lice had high levels of population isolation consistent with a paucity of horizontal transmission even at the very fine geographic scale of a single alfalfa field. We also found marked genetic discontinuity in louse populations corresponding with host subspecies and little, if any, admixture in the louse genetic groups even though the lice are closely related. The correlation of louse infrapopulation differentiation with host interaction at multiple scales, including across a discontinuity in pocket‐gopher habitat, suggests that host behaviour is the primary driver of parasite genetics. This observation makes sense in light of the life histories of both chewing lice and pocket gophers and provides a powerful explanation for the well‐documented pattern of parallel cladogenesis in pocket gophers and chewing lice.     

Survivorship of penstemon grandiflorus in an oak woodland: combined effects of fire and pocket gophers

Summary The effects of fire and pocket gophers, Geomys bursarius, on the survivorship of Penstemon grandiflorus growing in an oak woodland in Minnesota were studied from 1986 to 1990. Plants growing in sparse vegetation experienced mortality rates twice that of plants growing in dense vegetation. This difference was due partly to pocket gophers whose earth moving activities reduce the density of vegetation and bury and kill individual Penstemon plants. Laboratory feeding trials showed that gophers readily eat Penstemon, particularly the fleshy roots. An experiment involving the removal of 25–75% of the root tissue in 90 plants showed that root loss significantly reduced survivorship, suggesting that gopher herbivory might also kill plants. When gophers were experimentally excluded, plants growing in sparse vegetation exhibited significantly lower mortality rates than those growing in dense vegetation. Plants in the smallest size class exhibited reduced survivorship following a late spring burn; however, overall patterns of survivorship of plants in burned areas did not differ markedly from those in the unburned areas. A longitudenal analysis of plants with different reproductive histories revealed no survivorship cost to reproduction. Mortality rates decreased with increasing plant size. Small plants were more likely to be killed by fire and by being buried under gopher mounds. Differences in underground energy reserves of small and large plants can account for most of the survivorship patterns observed in this study. The study shows that within openings of the oak woodland, fire and gophers reduce the survival of individual Penstemon plants. Nevertheless, since both gophers and fire also serve to perpetuate suitable habitat in the woodland, Penstemon is ultimately dependent on both for its long term persistence in the landscape.     

Influence of pocket gopher mounds on a Texas coastal prairie

Summary Effects of pocket gopher (Geomys attwateri) mound-building activity on plant community composition and soil nutrient concentrations were investigated in south Texas on both burned and unburned coastal prairie sites. Pocket gophers deposited large amounts of soil which were lower in nutrient content than randomly-collected samples. Above-ground plant biomass was greater around mounds than in random samples mainly because of increased dicots around mounds on the burned area when compared with random samples on the same area. Pocket gophers may have concentrated their activities (and therefore, mounds) in areas with higher dicot biomass on the burned area since they prefer perennial dicots as food, or the presence of mounds may have ameliorated the apparent negative effect of fire on dicots.     

Phylogeny of Geomydoecus and Thomomydoecus pocket gopher lice (Phthiraptera: Trichodectidae) inferred from cladistic analysis of adult and first instar morphology

The phylogeny for all 122 species and subspecies of chewing lice of the genera Geomydoecus and Thomomydoecus (Phthiraptera: Trichodectidae) hosted by pocket gophers (Rodentia: Geomyidae) is estimated by a cladistic analysis of fifty-eight morphological characters obtained from adults and first instars. The data set has considerable homoplasy, but still contains phylogenetic information. The phylogeny obtained is moderately resolved and, with some notable exceptions, supports the species complexes proposed by Hellenthal and Price over the the last two decades. The subgenera G. (Thaelerius) and T. (Thomomydoecus) are both shown to be monophyletic, but the monophly of subgenus T. (Jamespattonius) could not be confirmed, perhaps due to the lack of first-instar data for one of its component species. The nominate subgenus of Geomydoecus may be monophyletic, but our cladogram was insufficiently resolved to corroborate this. Mapping the pocket gopher hosts onto the phylogeny reveals a consistent pattern of louse clades being restricted to particular genera or subgenera of gophers, but the history of the host-parasite association appears complex and will require considerable effort to resolve.     

Comparative body size relationships in pocket gophers and their chewing lice

In this paper, we use the method of independent contrasts to study body size relationships between pocket gophers and their chewing lice, a host-parasite system in which both host and parasite phylogcnies are well studied. The evolution of body size of chewing lice appears to be dependent only on the body size of their hosts, which confirms the 1991 findings of Harvey and Keymer. We show that there is a positive relationship between body size and hair-shaft diameter in pocket gophers, and that there is also a positive relationship between body size and head-groove width in chewing lice. Finally, we show a positive relationship between gopher hair-shaft diameter and louse head-groove width. We postulate that changes in body size of chewing lice are driven by a mechanical relationship between the parasite's head-groove dimension and the diameter of the hairs of its host. Louse species living"on larger host species may be larger simply because their hosts have thicker hairs, which requires that the lice have a wider head groove. Our study of gopher hair-shaft diameter and louse head-groove dimensions suggest that there is a 'lock-and-key' relationship between these two anatomical features.