Walter G. Whitford

Biological Feedbacks in Global Desertification

Studies of ecosystem processes on the Jornada Experimental Range in southern New Mexico suggest that longterm grazing of semiarid grasslands leads to an increase in the spatial and temporal heterogeneity of water, nitrogen, and other soil resources. Heterogeneity of soil resources promotes invasion by desert shrubs, which leads to a further localization of soil resources under shrub canopies. In the barren area between shrubs, soil fertility is lost by erosion and gaseous emissions. This positive feedback leads to the desertification of formerly productive land in southern New Mexico and in other regions, such as the Sahel. Future desertification is likely to be exacerbated by global climate warming and to cause significant changes in global biogeochemical cycles.

Impacts of shrub encroachment on ecosystem structure and functioning: towards a global synthesis

Encroachment of woody plants into grasslands has generated considerable interest among ecologists. Syntheses of encroachment effects on ecosystem processes have been limited in extent and confined largely to pastoral land uses or particular geographical regions. We used univariate analyses, meta-analysis and structural equation modelling to test the propositions that (1) shrub encroachment does not necessarily lead to declines in ecosystem functions and (2) shrub traits influence the functional outcome of encroachment. Analyses of 43 ecosystem attributes from 244 case studies worldwide showed that some attributes consistently increased with encroachment (e.g. soil C, N), and others declined (e.g. grass cover, pH), but most exhibited variable responses. Traits of shrubs were associated with significant, though weak, structural and functional outcomes of encroachment. Our review revealed that encroachment had mixed effects on ecosystem structure and functioning at global scales, and that shrub traits influence the functional outcome of encroachment. Thus, a simple designation of encroachment as a process leading to functionally, structurally or contextually degraded ecosystems is not supported by a critical analysis of existing literature. Our results highlight that the commonly established link between shrub encroachment and degradation is not universal.

The Role of Mites and Nematodes in Early Stages of Buried Litter Decomposition in a Desert

We studied changes in populations of mites, nematodes, bacteria, and fungi in buried creosote bush litter treated with selected inhibitors. Elimination of microarthropods (primarily tydeid mites) resulted in increased numbers of bacteriophagic nematodes and reduction in numbers of bac- teria; elimination of both nematodes and microarthropods resulted in increased numbers of bacteria compared to untreated controls. Fungal grazing mites, Pyemotidae, and fungivorous nematodes, Aphelenchus sp., increased in numbers between days 25 and 30, reducing the fungi on untreated leaves but not on stems and petioles, while mean length of fungal hyphae increased in insecticide- treated leaves. Elimination of mites resulted in a 40% reduction in decomposition suggesting that in a desert, tydeid mites affect decomposition of buried litter by regulating the population size of the bacterial grazers, cephalobid nematodes. In our previous study (Santos and Whitford 1981), we found that tydeid mites were the initial arthropod colonizers of buried creosote bush litter and that or- ganic matter loss was significantly reduced in the ab- sence of these mites. We hypothesized that the tydeid mites were predators on free-living nematodes, and regulated decomposition by regulating the population size of microbial grazers (the nematodes).

Chihuahuan Desert annuals: importance of water and nitrogen

We examined the effects of water supplementation and nitrogen amendment on biomass, cover, and density of annual plants on a termite-free and a termite-present area in the Chihuahuan Desert. Soil moisture was higher in the termite than in the termite-free plots, and in the watered than in the unwatered plots during the spring and summer. There were no differences in soil moisture among plots during the winter. Soil nitrogen was higher in the termite-free than in the termite plots. There were no differences in total plant biomass produced in termite and termite-free areas. There were significant differences in relative abundances of species among treatments. natural rainfall was sufficient for maximum spring-annual biomass development on all plots except for the termite-free unfertilized, unwatered plots. These were the driest plots but had high soil nitrogen. Most of the herbaceous species responded to the water amendments by lengthening growing seasons, increasing density, or increasing biomass. When there was sufficient water for most of the spring annuals, high soil nitrogen levels favored increased densities and biomasses of Descurainia pinnata and Lepidium lasiocarpum. The absence of C4 summer annuals in the high-nitrogen plots suggests that relatively high soil nitrogen adversely affected the summer annuals. Termite-free watered plots had higher soil moisture than the termite-unwatered plots, but summer annuals were relatively abundant on the latter. Water amendments had a greater effect on the species abundances in the termite-free area than in the one with termites. In the area with termites, nitrogen amendments had a greater effect on species abundances. Species diversity and richness were affected by fertility as was species composition. This study demonstrates that we must understand patterns of soil nitrogen availability and processes affecting nitrogen availability in addition to water availability, in order to understand productivity and species composition of Chihuahuan Desert annual plants.

The influence of subterranean termites on the hydrological characteristics of a Chihuahuan desert ecosystem

SummaryRainfall simulation at an average intensity of 124 mm·h-1 was used to compare infiltration and run off on arid areas where subterranean termites had been eliminated four years prior to the initiation of the study (termite free) with adjacent areas populated by subterranean termites (termites present). Infiltration rates on termite free plots with less than 5% perennial plant cover were significantly lower 51.3±6.8 mm·h-1 than rates on comparable termites present plots 88.4±5.6 mm·h-1. On plots centered on Larrea tridentata shrubs, there were no differences in infiltration rates with or without termites. Plots with shrub cover had the highest infiltration rates 101±6 mm·h-1. Highest run-off volumes were recorded from termite free <5% grass cover plots and the lowest from plots with shrubs. There were no differences in suspended sediment concentrations from termites present and termite free plots. Average bed load concentration was more than three times greater from termite free, <5% cover plots than from termites present, <5% cover plots.The reduction in infiltration, high run-off volumes and high bedloads from termite free areas without shrub cover is related to increased soil bulk density resulting from the collapse of subterranean galleries of the termites that provide avenues of bulk flow into the soil. Subterranean termites affect the hydrology of Chihuahuan desert systems by enhancing water infiltration and retention of top soil. The presence of a shrub canopy and litter layer cancels any effect of subterranean termites on hydrological parameters. Since approximately 2/3 of the area is not under shrub canopies, subterranean termites are considered to be essential for the maintenance of the soil water characteristics that support the present vegetation.

Field soil aggregate stability kit for soil quality and rangeland health evaluations

Abstract Soil aggregate stability is widely recognized as a key indicator of soil quality and rangeland health. However, few standard methods exist for quantifying soil stability in the field. A stability kit is described which can be inexpensively and easily assembled with minimal tools. It permits up to 18 samples to be evaluated in less than 10 min and eliminates the need for transportation, minimizing damage to soil structure. The kit consists of two 21×10.5×3.5 cm plastic boxes divided into eighteen 3.5×3.5 cm sections, eighteen 2.5-cm diameter sieves with 1.5-mm distance openings and a small spatula used for soil sampling. Soil samples are rated on a scale from one to six based on a combination of ocular observations of slaking during the first 5 min following immersion in distilled water, and the percent remaining on a 1.5-mm sieve after five dipping cycles at the end of the 5-min period. A laboratory comparison yielded a correlation between the stability class and percent aggregate stability based on oven dry weight remaining after treatment using a mechanical sieve. We have applied the method in a wide variety of agricultural and natural ecosystems throughout western North America, including northern Mexico, and have found that it is highly sensitive to differences in management and plant community composition. Although the field kit cannot replace the careful laboratory-based measurements of soil aggregate stability, it can clearly provide valuable information when these more intensive procedures are not possible.

Water and nitrogen effects on growth and allocation patterns of creosotebush in the northern Chihuahuan Desert.

A field experiment udng 2 patterns of irrigation and 1 level of nitrogen fertilizer (10 g-N m-2) was conducted in order to discern water and nitrogen interactions that may control production of creosotebush, (Laweu tridentatu (D.C.) Cov. The 2 patterns of irrigation simulated precipitation from small, frequent events (6 mm water added weekly) or large, infrequent events (25 mm water added monthly). Understanding the factors controlling the production of this rangeland shrub may aid in the development of strategies for its management. Vegetative growth occurred mostly during March-May (spring)

The effect of water and nitrogen amendments on photosynthesis, leaf demography, and resource-use efficiency in Larrea tridentata, a desert evergreen shrub

SummaryIn the Chihuahuan Desert of southern New Mexico, both water and nitrogen limit the primary productivity of Larrea tridentata, a xerophytic evergreen shrub. Net photosynthesis was positively correlated to leaf N, but only in plants that received supplemental water. Nutrient-use efficiency, defined as photosynthetic carbon gain per unit N invested in leaf tissue, declined with increasing leaf N. However, water-use efficiency, defined as the ratio of photosynthesis to transpiration, increased with increasing leaf N, and thus these two measures of resource-use efficiency were inversely correlated. Resorption efficiency was not significantly altered over the nutrient gradient, nor was it affected by irrigation treatments. Leaf longevity decreased significantly with fertilization although the absolute magnitude of this decrease was fairly small, in part due to a large background of insect-induced mortality. Age-specific gas exchange measurements support the hypothesis that leaf aging represents a redistribution of resources, rather than actual deterioration or declining resource-use efficiency.

Foraging in Seed-Harvester Ants Progonomyrmex SPP

Pogonomyrmex rugosus, a group forager, foraged preferentially on seeds of plant species which shed large quantities of seeds at I time. Pogonomyrmex desertorum, an individual forager took seeds in relation to their availability but concentrated on grass seeds late in the growing season. Pogonomyrmex californicus selected seed species which allowed it to avoid contact with its congeners. During a dry year, P. rugosus exhibited little selectivity, supporting the idea that selectivity should be decreased under conditions of low food abundance. Pogonomyrmex spp. removed a sig- nificant fraction of the seed production of only 1 species, Boutleoua barbata. It is suggested that Pogonomyrmex foraging may affect the relative abundance of plant species.

The importance of the biodiversity of soil biota in arid ecosystems

The importance of soil biota in maintaining ecosytem integrity is examined by a review of studies of soil processes and soil biota in arid ecosystems. In decomposition and mineralization processes, there is a temporal succession of microarthropod and nematode species. Tydeid mites are keystone species in the early stages of decomposition. Soil pore neck size variation affects the spatial distribution of soil biota and requires that species differ in size to provide efficient processing of organic matter. In arid ecosystems, the sub-set of soil biota that is active at any point in time is determined by the soil water potential and soil temperature. The structure of soil microarthropod assemblages is similar in several arid regions of the world and abundance and diversity are directly related to quantity of litter accumulations and soil organic matter. The unique life histories and behavioural characteristics of desert soil macrofauna (termites and ants) determine the effects of these organisms on soil properties and soil formation. The soil biota, by affecting the spatial and temporal distribution of essential sesources (water and nutrients), are essential to the maintenance of the ecosystem integrity of arid ecosystems.