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.

Mechanisms for the keystone status of kangaroo rats: graminivory rather than granivory?

Graminivory by kangaroo rats (Dipodomys spp.) was investigated as a potential mechanism for the keystone role of these rodents in the dynamics of desert grasslands. Experiments confirmed that Ord's kangaroo rats (Dipodomys ordii) cut and consumed a large proportion of the tillers of three Chihuahuan Desert tussock-forming grass species. Field observations indicated that the characteristically cut grass tillers were absent from all-rodent and medium-sized kangaroo rat exclosures, but were frequent in large-sized kangaroo rat and rabbit exclosures, indicating that the medium-sized kangaroo rats (D. ordii, D. merriami) were responsible for grass cutting. Tiller waste as a percentage of peak standing crop ranged from 7% in grassland habitats to 0.7% in Flourensia cernua shrubland. Of the 13 species of perennial, tussock-forming grasses measured, only one, Muhlenbergia porteri, had no tillers cut by kangaroo rats. This study demonstrates that the keystone role of kangaroo rats in Chihuahuan Desert grassland ecosystems is probably the result of their graminivory.

Pollination ecology of Yucca elata : An experimental study of a mutualistic association.

The pollination biology of a population of 250 Yucca elata (Liliaceae) plants was studied in southern New Mexico. Yucca elata and the prodoxid yucca moth Tegeticula yuccasella have a mutualistic association that is essential for the successful sexual reproduction of both species. However, a wide range of other invertebrate species visit flowers during the day and at night. Our aim was to quantify the role of yucca moths and other invertebrate visitors in pollination and fruit set, using manipulative field experiments. Inflorescences were bagged during the day or night (N=12 inflorescences) to restrict flower visitors to either nocturnal or diurnal groups. Yucca moths were active exclusively nocturnally during the flowering period and thus did not visit inflorescences that were unbagged during the day. None of the 4022 flowers exposed only to diurnal visitors set fruit, whereas 4.6% of the 4974 flowers exposed only to nocturnal visitors (including yucca moths) produced mature fruit. The proportion of flowers producing fruit in the latter treatment was not significantly different from unbagged control inflorescences. In a series of experimental manipulations we also determined that: (1) flowers opened at dusk and were open for two days on average, but were only receptive to pollen on the first night of opening; (2) pollen must be pushed down the stigmatic tube to affect pollination; and (3) most plants require out-cross pollination to produce fruit. The combination of these results strongly suggests that yucca moths are the only species affecting pollination in Y. elata, and that if another species was to affect pollination, it would be a rare event.

Phytophagous insects enhance nitrogen flux in a desert creosotebush community.

We tested the hypothesis that herbivorous insects on desert shrubs contribute to short-term nitrogen cycling, and increase rates of nitrogen flux from nutrient rich plants. Creosotebush (Larrea tridentata) shrubs were treated with different combinations of fertilizer and water augmentations, resulting in different levels of foliage production and foliar nitrogen contents. Foliage arthropod populations, and nitrogen in canopy dry throughfall, wet throughfall and stemflow were measured to assess nitrogen flux rates relative to arthropod abundances on manipulated and unmanipulated shrubs over a one-month period during peak productivity. Numbers and biomass of foliage arthropods were significantly higher on fertilized shrubs. Sap-sucking phytophagous insects accounted for the greatest numbers of foliage arthropods, but leaf-chewing phytophagous insects represented the greatest biomass of foliage arthropods. Measured amounts of bulk frass (from leaf-chewing insects) were not significantly different among the various treatments. Amounts of nitrogen from dry and wet throughfall and stemflow were significantly greater under fertilized shrubs due to fine frass input from sap-sucking insects. Increased numbers and biomass of phytophagous insects on fertilized shrubs increased canopy to soil nitrogen flux due to increased levels of herbivory and excrement. Nitrogen excreted by foliage arthropods accounted for about 20% of the total one month canopy to soil nitrogen flux, while leaf litter accounted for about 80%.

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

In 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.

Interplant variation in creosotebush foliage characteristics and canopy arthropods.

We conducted a field study to test the hypothesis that creosotebush (Larrea tridentata) shrubs growing in naturally nutrient-rich sites had better quality foliage and supported greater populations of foliage arthropods than shrubs growing in nutrient-poor sites. Nutrient-rich sites had significantly higher concentrations of soil nitrogen than nutrient-poor sites. Multivariate analysis of variance revealed significant differences between high nutrient and low nutrient shrubs based on a number of structural and chemical characteristics measured. High nutrient shrubs were larger, had denser foliage, greater foliage production, higher concentrations of foliar nitrogen and water, and lower concentrations of foliar resin than low nutrient shurbs. Numbers of foliage arthropods, particularly herbivores and predators, were significantly higher on high nutrient shrubs. Shrub characteristics and foliage arthropod abundances varied considerably from shrub to shrub. Shrub characteristics representing shrub size, foliage density, foliage growth, and foliar nitrogen and water concentrations were positively correlated with arthropod abundances. Foliar resin concentrations were negatively correlated with foliage arthropod abundances. The positive relationship between creosotebush productivity and foliage arthropods is contradictory to the tenet that physiologically stressed plants provide better quality foliage to insect herbivores.

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.

A conceptual model for primary productivity, decomposition and nitrogen cycling in the Chihuahuan creosotebush desert.

The conceptual framework for a simulation model of primary productivity, decomposition and nitrogen cycling in a shrub-dominated desert ecosystem in southern New Mexico is presented. This model is based on our previous attempt to simulate carbon allocation patterns in the desert shrub Larrea tridentata Cov., which demonstrated that moisture patterns alone are insufficient to predict desert productivity. These results, as well as others, suggest that mineral nutrients, especially nitrogen, may also be an important determinant of productivity in arid environments. Our current research in the Chihuahuan desert is directed towards elucidating the numerous biotic and abiotic interactions that determine the rates and directions of carbon, nitrogen and water fluxes in this ecosystem. The development of this working model will serve as a tool to accomplish three major objectives: (1) to synthesize the large amount of existing data on decomposition and nitrogen cycling in deserts, (2) to quantify our present state of knowledge about the structure and function of ecosystem components important in carbon and nitrogen dynamics in deserts, and (3) to address hypotheses concerning the complex mechanisms of interactions and feedbacks among the organisms involved in carbon and nitrogen exchanges in deserts.

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

Rainfall 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.

The relative contributions of termites and microarthropods to fluff grass litter disappearance in the Chihuahuan Desert.

We tested the hypotheses that both subterranean termites and soil microarthropods are important in the disappearance of fluff grass (Erioneuron pulchellum) litter on the soil surface by an experiment designed to separate termite and microarthropod effects. Subterranean termites (Gnathamitermes tubiformans) removed more than 50% of the fluff grass litter in one year.Since there was no difference in mass loss of fluff grass with microarthropods present or excluded, they had no effect on decomposition of fluff grass litter. Microarthropod densities increased during the first 3 months then slowly decreased. The densities of microarthropods in fluff grass litter were too low to have a measurable effect on decomposition even if we assumed that the microarthropods consumed litter equivalent to their body weight each day.

The failure of nitrogen and lignin control of decomposition in a North American desert.

We measured mass losses of both buried and surface litter of six litter types: leaves of the perennial evergreen shrub, Larrea tridentata, leaves of the winter deciduous perennials Fluorensia cernua, Prosopis glandulosa and Chilopsis linearis (a desert riparian species), an evergreen monocot, Yucca elata, and a mixture of annual plants. These species differed in lignin content and carbon-nitrogen ratios. There was no correlation between rates of mass loss and percent lignin, carbon-nitrogen ratio, or lignin-nitrogen ratio. The leaves of F. cernua and the mixed annuals exhibited the highest rates of mass loss. Surface litter of Y. elata, the mixed annuals and C. linearis exhibited higher mass loss than buried litter of the same species. The patterns of mass loss of buried and surface litter differed with buried litter mass loss occurring as a negative exponential and surface litter exhibiting low rates in winter and spring and high rates in summer. There was no correlation between mass loss in surface bags that were field exposed for 1 month and actual evapotranspiration (AET) but there was a correlation between AET and mass losses in buried litter. A model relating mass loss to AET and initial lignin content underestimated mass losses in all species examined.

Contributions of subterranean termites to the "economy" of Chihuahuan desert ecosystems.

We examined the role of subterranean termites in decomposition of cattle dung, various herbaceous plant species and wood in a Chihuahuan desert ecosystem. From July-September, termites removed dung at a rate of 0.63 g day-1 accounting for a percent mass loss of 19.5-100%. During the autumn subterranean termites consumed more than 50% of the leaves of the shrub Larrea tridentata, the grass Erioneuron pulchellum and annual plant Lepidium lasiocarpum and Baileya multiradiata but used very little of two other annuals Eriastrum diffusum and Eriogonum trichopes. Yucca inflorescence stalks on plots with termites lost 23% of their original mass in 30 months while those on termite free soils lost 11%. Elimination of termites resulted in reduction of fluff grass, Erioneuron pulchellum biomass, thereby affecting the structure of the ecosystem.

The role of microarthropods and nematodes in decomposition in a semi-arid ecosystem.

We sampled the soil microarthropod community monthly in the oak-mesquite sand hill ecosystem. Small fungiphagous prostigmated mites (pyemotids, lordalychids and tarsonemids) that dominated the soil fauna in winter were replaced by large predaceous mites (rhodacarids and laelapids) in summer and autumn.We compared organic matter loss and microarthropod and nematode populations in shinnery oak (Quercus harvardii) using insecticide and untreated litter in fiberglass litterbags.Microarthropods extracted from litterbags showed a seasonal pattern similar to the soil cores except that collembolans and psocopterans were abundant in the litter and not in the soil cores. Numbers of free living nematodes were consistently greater than from untreated litter. The ratio of non-stylet to stylet bearing nematodes extracted from litter decreased from 4:1 in one month bags to 0.8:1.0 in the one year bags. Laboratory experiments showed that rhodacarid mites fed voraciously on nematodes.Untreated litter exhibited higher rates of organic matter loss than the insecticide treated litter; 20% and 35% respectively.We suggest that the abundant mesostigmatid mites prey on free living nematodes and that eliminating the predators allows the nematodes to overgraze the fungi and bacteria. The soil modifies the microclimate in buried litter allowing for higher biological activity, hence higher rates of decomposition.

The relationship between wood litter infall and relative abundance and feeding activity of subterranean termites Reticulitermes spp. in three southeastern coastal plain habitats.

We estimated the densities of termites in dead wood using a point-quarter technique in four habitats: pine plantations subject to control burning and unburned, lowland hardwood forest, and turkey oak woodland. The burned pine plantation had the highest estimated termite density, 13×106·ha-1 unburned pine plantation and lowland hardwood had estimated densities of 2.6×106·ha-1 and 2.2×106·ha-1, respectively, and the turkey oak woodland had an estimated densities of 61.9×103·ha-1. There were varying percentages of Reticulitermes flavipes and R. virginicus in the various habitats. There was nearly linear increase in percentage of pine blocks attacked by termites in the pine and hardwood forests and by the end of the growing season, nearly all had been channelized by termites. In the turkey oak habitat 70% of the pine blocks were channelized. Termites removed between 3% and 12% of the original mass of over one-fourth of the pine blocks during the growing season.Wood litter in fall was highest in the lowland hardwood forest: 2869 kg·ha-1·yr-1. Wood litter input in the long leaf pine plantation, 792 kg·ha-1·yr-1 was nearly equivalent to the three year average wood litter in fall in a Danish oak forest.

Nutrient cycling by the subterranean termite Gnathamitermes tubiformans in a Chihuahuan desert ecosystem.

We estimated the density of subterranean termites Gnathamitermes tubiformans at 800,000 · ha-1 for a standing crop biomass of 2 kg · ha-1 Predation losses were estimated to be 5,73 kg · ha-1 · yr-1 representing the major release of nutrients from termites to surficial soil layers. Nutrient fluxes from termites to predators amounted to 410g N·ha-1·yr-1, 33 g S · ha-1 · yr-1 and 19 g P · ha-1 · yr-1. These fluxes account for 8% of the litter N, 1.5% of the litter P and 2.9% of the litter S. The termites fixed an estimated 66 g · ha-1 · yr-1 atmospheric N and returned an estimated 100 g · ha-1 · yr-1 in the surface gallery carton. Since losses of elements from subterannean termites were greater than standing crops, we estimated an annual turnover of N at 3.5 times per year, P of 2.5 times per year, and S of 2.5 per times per year.Since surface foraging, predation and alate flights are pulse regulated by rainfall, nutrient flows through subterranean termites are episodic and releases of nutrients accumulated in termite biomass preceeds or is coincident with productivity "pulses" of some shallow rooted plants. We propose that subterranean termites are important as regulators in desert nutrient cycles.