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.     

Community and population dynamics of serpentine grassland annuals in relation to gopher disturbance

Summary This study examines the effects of soil disturbance by gophers on patterns of species abundance in an annual grassland community on serpentine soil. We assessed production, dispersal and storage of seed, germination, survivorship and growth of the most abundant species in undisturbed vegetation and on gopher mounds. Fewer seeds of the dominant species were dispersed onto gopher mounds due to the limited movement of seeds from within the closed vegetation. Species with taller flowering stalks were more likely to colonise gopher mounds. The timing of gopher disturbance in relation to the timing of seed fall determined which species could colonise mounds. Lower numbers of seeds falling onto gopher mounds resulted in lower seedling densities of several species compared with undisturbed areas. Survivorship of the commonest species differed between undisturbed areas and gopher mounds formed at different times of year. This resulted in characteristic spectra of species abundance on the different microhabitats, giving rise to distinct spatial patterning in the community. Plants growing on gopher mounds were generally larger and produced more seed than plants in undisturbed vegetation. We suggest that continued gopher disturbance is a factor allowing several species, including perennial grasses, to persist in this community.     

Vegetation associations of birds wintering in a California oak woodland

Conditions in wintering areas can have persistent effects throughout the year for bird populations. Most studies, however, are conducted during the breeding season and the non‐breeding ecology of many species remains poorly understood. We assessed vegetation associations of birds wintering in a diverse, well‐structured oak woodland in coastal‐central California. We calculated density estimates for 20 bird species and examined correlations between bird densities and vegetation characteristics as described by a principal components analysis. Ruby‐crowned Kinglets (Regulus calendula) and Hutton's Vireos (Vireo huttoni) were positively associated with vegetation characteristic of dense, floristically diverse woodland with abundant coast live oak (Quercus agrifolia) cover and a structurally complex understory. Conversely, White‐breasted Nuthatches (Sitta carolinensis) and Western Bluebirds (Sialia mexicana) were negatively correlated with dense woodland and occurred at the highest densities on plots with large widely spaced blue oaks (Q. douglasii) and a sparse understory. Thirteen of the remaining 16 species were more abundant in dense woodland patches, but these relationships were not statistically significant. Our results are in agreement with those of previous studies and suggest that evergreen oak trees and a dense understory provide key resources (e.g., cover and food) for some bird species during the winter. However, areas with large widely spaced deciduous oaks are important habitats for other species. We therefore suggest that dense patches of live oaks and shrubs should be retained, but a mosaic of dense and sparse woodland is necessary to accommodate the entire suite of oak‐woodland birds.     

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.     

Spatial variation of seedling distribution in an east Mediterranean pine woodland at the beginning of post-fire succession

Most of the area in pine woodlands is occupied by perennial seeders that regenerate from seeds in the first winter after the fire and by annuals. Control of the germination in the regenerating vegetation after wildfire is therefore a primary ecological component of the post-fire succession in this ecosystem. The aim of the study presented here was to determine the distribution of Pinus, Cistus and other plants seeds around burned Pinus halepensis trees, and to measure the conditions related to seed germination in the upper soil layers in the same locations. The study was carried out in a 50-year old planted Pinus halepensis woodland that was burned down by a wildfire in July 1995. The variation of seedbank density was determined by collecting samples under the canopies of burned trees and in a nearby open area. Pine seedbank density decreased and that of Cistus and annuals increased with increasing distance from the burned trunks. Most pine seeds were present in the ash layer while those of the other plants were in the soil. In situ germination experiments showed that seedling density decreased with distance from the burned trunks while the proportion of pines in the seedling population increased. This was a result of seedbank variation and germination inhibition by the high pH conditions caused by the ash. The establishment of sparse pine seedling under the dead tree canopies insured their rapid development without interference by other plants and played a key role in the regeneration and stability of the pine woodland community. The concomitant mass germination of the perennial seeders in the rest of the area prevented invasion by annuals.     

Edaphic limitations to growth and photosynthesis in Sierran and Great Basin vegetation

Summary Soils derived from hydrothermally altered andesite support unique communities of Sierran conifers (Pinus ponderosa Laws. and P. jeffreyi Grev. and Balf.) amongst sagebrush (Artemisia tridentata Nutt.) vegetation in the western Great Basin. Plants grown in soil derived from hydrothermally altered bedrock had lower growth rates, total biomass, and net photosynthetic rates than plants grown in soil derived from unaltered andesite of the same formation. Total dry mass was 10 to 28% lower for conifers grown in altered soil whereas dry mass of Artemisia tridentata and Bromus tectorum L. was reduced by over 90%. Results from a nutrient amendment experiment indicated that low phosphorus was the dominant limitation in altered soil, and phosphorus-deficiency affected growth primarily by limiting leaf area development rather than direct inhibition of photosynthesis. The proportionately greater reduction of biomass for Artemisia and Bromus grown in altered soil supports our hypothesis that Great Basin vegetation is excluded from altered soil by intolerance to nutrient deficiency. The Sierran conifers growing on this rock type are therefore free of competition for water with Great Basin vegetation and are able to persist in an exceptionally dry climate.     

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.     

Survivorship patterns in an annual plant community

Summary Age-specific survivorship was studied in a number of plant species which occur in a distinct, predominantly annual plant community of Texas, in association with the activities of pocket gophers. Of eleven species analyzed in detail, all functioned as annuals. Contrary to theoretical expectations, the species had low rates of juvenile mortality, and did not exhibit the demographic specializations of weedy or colonizing species. It is suggested that the annual habit in this case may be adaptive in sites with continual disturbance, such as that due to pocket gopher activity.     

Habitat associations of California ground squirrels and Botta's pocket gophers on levees in California

Vegetation management on levees, especially removal of trees and shrubs, might affect burrowing mammals that are considered threats to levee integrity. We evaluated habitat associations of California ground squirrels (Otospermophilus beecheyi) and Botta's pocket gophers (Thomomys bottae) on levees in the Sacramento Valley to assess the effects of levee vegetation management on these species. Using burrows as an indirect measure of mammal presence, we found that the presence of trees had a negative effect on the occurrence and abundance of ground squirrels on levees, and on the location of their burrowing activities on the levee slope, potentially because visual occlusion caused by tall woody vegetation impedes detection of predators. Similarly, trees had a negative effect on the abundance of pocket gophers on levees and on the location of their burrowing activities on the levee slope, probably because of the effect of tree cover on food availability. The conversion of woodland habitats to grasslands on levees most likely will result in increased occurrence, abundance, or both of ground squirrels and pocket gophers, and thereby increase the potential threat that their burrowing activities pose to levee integrity. © 2012 The Wildlife Society.     

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.     

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.     

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     

Soil translocation by pocket gophers in a Mima moundfield

Summary We measured soil translocation due to the tunneling of valley pocket gophers (Thomomys bottae) in a Mima moundfield at Miramar Mounds National Landmark, San Diego, California, from December, 1984 through December, 1985. We placed 1-l soil plugs containing 20 11-g iron pellets into pocket gopher tunnels at locations between mound tops and points about one mound radius beyond mound edges. After about 4–10 d, sites to which the marker-containing soil had been translocated were located with a metal detector and the horizontal and vertical displacements measured. Between 1 October and 15 May (the cooler, wetter portion of the year), pocket gophers removed an average of 63% of the experimental plugs and moved an average of 38% of the markers that we recovered. From 15 May through 1 October (the hotter, drier portion of the year), only 32% of plugs were cleared and 12% of the recovered markers were moved. On average, markers that were moved were displaced 41 cm moundward and 4.9 cm upward in elevation. The intensity of moundward translocation increased with distance from the mound center. At a distance of 0.5–1.0 mound radius beyond the edge of the mound, the moundward translocation tendency averaged 71 cm. The intensity of moundward translocation was also inversely related to maximum mound height. These observations provide strong support for the fossorial rodent hypothesis of Mima mound origin, and constitute a first step in development of a mathematical model of mound formation.     

On the Holocene vegetation history of the Argive Plain (Peloponnese,southern Greece)

Palynological investigations have been carried out on a sediment core from ancient Lake Lerna, a former fresh water lagoon in the western part of the Argive plain, Peloponnese, southern Greece. The sequence starts at 6800 B.P. (5700 cal B.C.). The lowest part of the pollen diagram shows a period of open deciduous oak woods, which may have been influenced by human impact already (Zone I). It is followed by a period of dense deciduous oak woods (Zone II), which lasted until the beginning of the Bronze Age ca. 4800 B.P. (3500 cal B.C.). Later, the diagram indicates strong human influence such as woodland clearance, the spread of maquis, phrygana and pine in Zones IIIa-IV. During the Archaic, Geometric and Classical periods after ca. 2700 B.P. (800 cal B.C.) there is evidence of a phase of extensive olive farming (Zone IIIb). In the same zone, after a period of scattered finds, there is an almost continuous Juglans curve. Zone IV is characterised by high pine values. In Zones I-II the evidence of evergreen Mediterranean plants is surprisingly small. In times with no discernible human influence (Zone II), deciduous oaks dominate, with no evidence for a climax vegetation of the Oleo-Ceratonion alliance. Olea europaea is the only species of that alliance traceable by its pollen in the diagram, while Ceratonia pollen is totally absent.     

Importance of the gradient in photosynthetically active radiation in a vegetation stand for leaf nitrogen allocation in two monocotyledons

Carex acutiformis and Brachypodium pinnatum were grown with a uniform distribution of photosynthetic photon flux density (PFD) with height, and in a vertical PFD gradient similar to the PFD gradient in a leaf canopy. Distribution of organic leaf N and light-saturated rates of photosynthesis were determined. These parameters were also determined on plants growing in a natural vegetation stand. The effect of a PFD gradient was compared with the effect of a leaf canopy. In Brachypodium, plants growing in a vegetation stand had increasing leaf N with plant height. However, distribution of leaf N was not influenced by the PFD gradient treatment. The gradient of leaf N in plants growing in a leaf canopy was not due to differences within the long, mostly erect, leaves but to differences between leaves. In Carex, however, the PFD gradient caused a clear increase of leaf N with height in individual leaves and thus also in plants. The leaf N gradient was similar to that of plants growing in a leaf canopy. Leaf N distribution was not affected by nutrient availability in Carex. In most cases, photosynthesis was positively related to leaf N. Hence, lightsaturated rates of photosynthesis increased towards the top of the plants growing in leaf canopies in both species and, in Carex, also in the PFD gradient, thus contributing to increased N use efficiency for photosynthesis of the whole plant. It is concluded that in Carex the PFD gradient is the main environmental signal for leaf N allocation in response to shading in a leaf canopy, but one or more other signals must be involved in Brachypodium.     

The effects of phosphorus nutrition and soil pH on the growth of some ancient woodland indicator plants and their interaction with competitor species

The forest under-storey herbs Anemone nemorosa, Lamiastrum galeobdolon and Veronica montana are generally considered indicator species of old, broadleaved woodland sites where the soil fertility is often low. In a glasshouse bioassay, however, all three species not only showed large positive growth responses to supplied P concentrations (0–10mgL ^–1) solutions, but also tolerated high P concentrations (20–40mgL ^–1), well above those normally found in their natural habitat. Plants responded by raising the concentrations of P in their shoot and root tissues and increasing their biomass, resulting in an increased P uptake. A shade-tolerant competitor species, Urtica dioica, also grew vigorously across the full range of P concentrations, restricting the growth of the woodland species. This emphasises the difficulty of establishing semi-natural woodland vegetation in the presence of competitor species, for example in situations where new woodlands are planted on fertile ex-agricultural soils containing large residual concentrations of P. The influence of soil pH on the growth and nutrient relations of A. nemorosa, L. galeobdolon, V. montana, Poa trivialisandU. dioicawas determined in a separate experiment using an ex-arable soil as the growing medium with pH levels adjusted from 7.4 to 5.8 and 4.3 respectively. Acidifying the soil enhanced growth, but reduced the concentrations of N, P and K in the leaves of all three woodland species, probably due to dilution of these minerals in the increased dry matter production. The competitor species (P. trivialis and U. dioica) responded in similar manner to the woodland indicator species. These results suggest that manipulating soil pH as a means of facilitating the establishment of woodland indicator species in new farm woods is unlikely, in the short term, to be effective where competitor species are present.     

Spatial relationships between plant litter,gopher disturbance and vegetation at different stages of old‐field succession

Abstract. Fine‐scale spatial patterns of small mammal disturbances and local accumulation of plant litter were studied together with the spatial pattern of vegetation in different stages of old‐field succession at Cedar Creek Natural History Area, Minnesota, USA. Seven stands from one to 66 years old were sampled. Presence of living plant species, local soil disturbances by pocket gophers (Geomys bursarius) and plant litter accumulation were recorded in 10 cm × 10 cm contiguous microquadrats along elliptical transects. Spatial patterns and associations were analyzed using information theory models. Dominant grasses were spatially independent, while subordinate functional groups were strongly dependent on the existing patchwork of dominant species, plant litter and gopher disturbances. Litter had consistent negative associations with subordinate functional groups in all but the initial years. Gopher disturbances were abundant but had weak and variable associations with vegetation. These results suggest that gopher disturbance does not facilitate the colonization of native prairie species and that diversity can be improved by controlling litter accumulation in Minnesota old‐fields.     

Rodent acorn selection in a Mediterranean oak landscape

Quercus suber, Quercus ilex and Quercus coccifera (Cork, Holm and Kermes oaks, respectively) are common evergreen oak species that coexist in the landscapes of the western part of the Mediterranean basin. Rodents are the main acorn predators and thus one of the main factors for understanding recruitment patterns in oaks. In this paper we analyse to what extent mice prefer acorns from one oak species over another in three oak species studied using acorn removal experiments and video tape recordings. Twenty labelled acorns from each of the three Quercus species (60 acorns) were placed in 40 cm×40 cm quadrats on each plot. Because selection might vary as a result of the vegetation context, we performed the trials in the five main vegetation types within the study area (four replicates in each vegetation type) in order to control for habitat influences on rodent acorn preferences (a total of 20 plots). The removal of 1,200 acorns occurred within 68 days. Mice removed 98.7% of the acorns. Q. ilex acorns were preferred over Q. suber and Q. coccifera in all vegetation types except in pine forest, where no acorn preferences were detected. Acorn removal rates differed with vegetation type, correlating positively with shrub cover. The distance at which acorns were displaced by rodents (mean =4.6 m±5.1 SD) did not differ between acorn species, but varied among vegetation types. Bigger acorns of Q. coccifera were selected only after Q. ilex and Q. suber acorns were depleted, while no size selection was detected for the latter two species. Thus, we conclude that rodents show preference for some oak acorns and that landscape context contributes significantly to rodent activities and decisions.     

Variation in species richness within and between calcareous (alvar) grassland stands: the role of core and satellite species

Anoda cristata is a widelydistributed annual weed in Mexico, which grows as agrestal or naturally indisturbed and undisturbed vegetation, respectively. Plants of this species aretolerated in orchards and corn-bean fields by Mexican ethnic groups ofCentral Mexico. Leaves of the plants are used as a food source, andoccasionallyseeds are sown in orchards. Because Mexicans have used Anodacristata for a long time, it is possible that ecological andmorphological characteristics of managed populations differ from those of wildpopulations. In this study, we analyzed phenotypic responses of two populationsof A. cristata (wild and agrestal) growing in twohabitats (forest and cultivated field) comparing survivorship, andlife history traits. Natural progenies from wild and agrestal populations ofA. cristata were transplanted into a cultivated field andin the understory of a pine forest in a reciprocal transplant experiment.Results showed that the habitat of transplant and the origin of populations hadsignificant effect on all plant characters measured (growth, phenology, andbiomass allocation), but the habitat x origin interaction wasnon-significant. In general, plants from the agrestal population grewfaster, reproduced earlier, and allocated more biomass to reproduction, thanplants from the wild population. Similarly, significant effects of habitat andorigin on plant survivorship were detected, but the origin of populationsexplained the largest proportion of variance in plant survival. Most traitswerephenotypically plastic but there were no differences in the magnitude ordirection of the response between populations. In contrast both populationsshowed differentiation for most character mean values. Populationdifferentiation is possibly the result of genetic differences driven byprocesses other than incipient domestication.     

Response to competition of bulbous geophyte Allium oleraceum differing in ploidy level

Experimental studies that explore the possible causes of ploidy distributions and niche differentiation are rare. Increased competitive ability may be advantageous for survival in dense vegetation and may strongly affect local and regional abundances of cytotypes and potentially contribute to invasion success. We compared survival, growth and reproduction of plants originating from bulbils of three cytotypes (2n = 4x, 5x, 6x) of Allium oleraceum growing with and without a competitor (Arrhenatherum elatius). There was a strong negative effect of competition but no effect of ploidy or ploidy × competition on survivorship, height and total dry mass of A. oleraceum, i.e. no support for different competitive abilities of the ploidy levels. However, slightly different responses of populations to competition treatments within all cytotypes suggest differentiation within cytotypes. Under competition, plant survivorship was low, surviving plants were small, had low dry mass and produced neither sexual nor asexual propagules. Without competition, plant survivorship was high, and cytotypes differed in three traits after 2 year's growth: dry mass of flowers, number of flowers and ratio of the dry mass of sexual to asexual propagules all decreased with increasing ploidy level. We additionally tested tetra‐ and pentaploids as to whether plants originating from different types of propagule (bulbils, seeds) differ in survivorship, growth and reproduction when growing with and without a competitor. Plants originating from bulbils had higher survivorship, were more robust, flowered earlier and produced more propagules when compared to plants originating from seeds and grown without competition. Under competition, differences in performance between plants originating from seeds and bulbils mostly disappeared, with higher survivorship only for plants originating from bulbils.