Mechanisms of grass response in grasslands and shrublands during dry or wet periods.

Multi-year climatic periods are expected to increase with global change, yet long-term data are often insufficient to document factors leading to ecological responses. We used a suite of long-term datasets (1993-2010) to examine the processes underlying different relationships between aboveground net primary production (ANPP) and precipitation in wet and dry rainfall periods in shrublands and grasslands in the Chihuahuan Desert. We hypothesized that trends in ANPP can be explained by different processes associated with their dominant grasses [Bouteloua eriopoda (grasslands); Sporobolus flexuosus (shrublands)] and with ecosystem properties that influence soil water dynamics with feedbacks to ANPP. We compared datasets on recruitment and growth for 7 years with no trend in precipitation followed by a 4-year drought and 5 consecutive wet years. We integrated these data in a simulation model to examine the importance of positive feedbacks. In grasslands, ANPP was linearly related to precipitation regardless of rainfall period, primarily as a result of stolon recruitment by B. eriopoda. A lag in responses suggests the importance of legacies associated with stolon density. In shrublands, ANPP was only related to rainfall in the wet period when it increased nonlinearly as the number of wet years increased. Seed availability increased in the first wet year, and seedling establishment occurred 2-4 years later. Increases in biomass, litter and simulated transpiration beginning in the third year corresponded with increases in ANPP. Understanding the processes underlying ecosystem dynamics in multi-year dry or wet periods is expected to improve predictions under directional increases or decreases in rainfall.

Hierarchical analysis of vegetation dynamics over 71 years: soil-rainfall interactions in a Chihuahuan Desert ecosystem.

Proliferation of woody plants in grasslands and savannas is a persistent problem globally. This widely observed shift from grass to shrub dominance in rangelands worldwide has been heterogeneous in space and time largely due to cross-scale interactions among soils, climate, and land-use history. Our objective was to use a hierarchical framework to evaluate the relationship between spatial patterns in soil properties and long-term shrub dynamics in the northern Chihuahuan Desert of New Mexico, USA. To meet this objective, shrub patch dynamics from 1937 to 2008 were characterized at patch and landscape scales using historical imagery and a recent digital soils map. Effects of annual precipitation on patch dynamics on two soils revealed strong correlations between shrub growth on deep sandy soils and above-average rainfall years (r = 0.671, P = 0.034) and shrub colonization and below-average rainfall years on shallow sandy soils (r = 0.705, P = 0.023). Patch-level analysis of demographic patterns revealed significant differences between shrub patches on deep and shallow sandy soils during periods of above- and below-average rainfall. Both deep and shallow sandy soils exhibited low shrub cover in 1937 (1.0% +/- 2.3% and 0.3% +/- 1.3%, respectively [mean +/- SD]) and were characterized by colonization or appearance of new patches until 1960. However, different demographic responses to the cessation of severe drought on the two soils and increased frequency of wet years after 1960 have resulted in very different endpoints. In 2008 a shrubland occupied the deep sandy soils with cover at 19.8% +/- 9.1%, while a shrub-dominated grassland occurred on the shallow sandy soils with cover at 9.3% +/- 7.2%. Present-day shrub vegetation constitutes a shifting mosaic marked by the coexistence of patches at different stages of development. Management implications of this long-term multi-scale assessment of vegetation dynamics support the notion that soil properties may constrain grassland remediation. Such efforts on sandy soils should be focused on sites characterized by near-surface water-holding capacity, as those lacking available water-holding capacity in the shallow root zone pose challenges to grass recovery and survival.

Use of low-temperature scanning electron microscopy to compare and characterize three classes of snow cover.

This study, which uses low-temperature scanning electron microscopy (LTSEM), systematically sampled and characterized snow crystals that were collected from three unique classes of snow cover: prairie, taiga, and alpine. These classes, which were defined in previous field studies, result from exposure to unique climatic variables relating to wind, precipitation, and air temperature. Snow samples were taken at 10 cm depth intervals from the walls of freshly excavated snow pits. The depth of the snow pits for the prairie, taiga, and alpine covers were 28, 81, and 110 cm, respectively. Visual examination revealed that the prairie snow cover consisted of two distinct layers whereas the taiga and alpine covers had four distinct layers. Visual measurements were able to establish the range of crystal sizes that occurred in each layer, the temperature within the pit, and the snow density. The LTSEM observations revealed the detailed structures of the types of crystals that occurred in the snow covers, and documented the metamorphosis that transpired in the descending layers. Briefly, the top layers from two of the snow covers consisted of freshly fallen snow crystals that could be readily distinguished as plates and columns (prairie) or graupel (taiga). Alternatively, the top layer in the alpine cover consisted of older dendritic crystal fragments that had undergone early metamorphosis, that is, they had lost their sharp edges and had begun to show signs of joining or bonding with neighboring crystals. A unique layer, known as sun crust, was found in the prairie snow cover; however, successive samplings from all three snow covers showed similar stages of metamorphism that led to the formation of depth hoar crystals. These changes included the gradual development of large, three-dimensional crystals having clearly defined flat faces, sharp edges, internal depressions, and facets. The study, which indicates that LTSEM can be used to enhance visual data by systematically characterizing snow crystals that are collected at remote locations, is important for understanding the physics of snowpacks and the metamorphosis that leads to potential avalanche situations. In addition, the metamorphosis of snow crystals must be considered when microwave radiometry is used to estimate the snow water equivalent in the winter snowpack, because large snow crystals more effectively scatter passive microwave radiation than small crystals.

Collecting, shipping, storing, and imaging snow crystals and ice grains with low-temperature scanning electron microscopy.

Methods to collect, transport, and store samples of snow and ice have been developed that enable detailed observations of these samples with a technique known as low-temperature scanning electron microscopy (LTSEM). This technique increases the resolution and ease with which samples of snow and ice can be observed, studied, and photographed. Samples are easily collected in the field and have been shipped to the electron microscopy laboratory by common air carrier from distances as far as 5,000 miles. Delicate specimens of snow crystals and ice grains survive the shipment procedures and have been stored for as long as 3 years without undergoing any structural changes. The samples are not subjected to the melting or sublimation artifacts. LTSEM allows individual crystals to be observed for several hours with no detectable changes. Furthermore, the instrument permits recording of photographs containing the parallax information necessary for three-dimensional imaging of the true shapes of snowflakes, snow crystals, snow clusters, ice grains, and interspersed air spaces. This study presents detailed descriptions of the procedures that have been used successfully in the field and the laboratory to collect, ship, store, and image snow crystals and ice grains.

Rime and graupel: description and characterization as revealed by low-temperature scanning electron microscopy.

Snow crystals, which form by vapor deposition, occasionally come in contact with supercooled cloud droplets during their formation and descent. When this occurs, the droplets adhere and freeze to the snow crystals in a process known as accretion. During the early stages of accretion, discrete snow crystals exhibiting frozen cloud droplets are referred to as rime. If this process continues, the snow crystal may become completely engulfed in frozen cloud droplets. The resulting particle is known as graupel. Light microscopic investigations have studied rime and graupel for nearly 100 years. However, the limiting resolution and depth of field associated with the light microscope have prevented detailed descriptions of the microscopic cloud droplets and the three-dimensional topography of the rime and graupel particles. This study uses low-temperature scanning electron microscopy to characterize the frozen precipitates that are commonly known as rime and graupel. Rime, consisting of frozen cloud droplets, is observed on all types of snow crystals including needles, columns, plates, and dendrites. The droplets, which vary in size from 10 to 100 microm, frequently accumulate along one face of a single snow crystal, but are found more randomly distributed on aggregations consisting of two or more snow crystals (snowflakes). The early stages of riming are characterized by the presence of frozen cloud droplets that appear as a layer of flattened hemispheres on the surface of the snow crystal. As this process continues, the cloud droplets appear more sinuous and elongate as they contact and freeze to the rimed crystals. The advanced stages of this process result in graupel, a particle 1 to 3 mm across, composed of hundreds of frozen cloud droplets interspersed with considerable air spaces; the original snow crystal is no longer discernible. This study increases our knowledge about the process and characteristics of riming and suggests that the initial appearance of the flattened hemispheres may result from impact of the leading face of the snow crystal with cloud droplets. The elongated and sinuous configurations of frozen cloud droplets that are encountered on the more advanced stages suggest that aerodynamic forces propel cloud droplets to the trailing face of the descending crystal where they make contact and freeze.

Irregular snow crystals: structural features as revealed by low temperature scanning electron microscopy.

For nearly 50 years, investigators using light microscopy have vaguely alluded to a unique type of snow crystal that has become known as an irregular snow crystal. However, the limited resolution and depth-of-field of the light microscope has prevented investigators from characterizing these crystals. In this study, a field-emission scanning electron microscope, equipped with a cold stage, was used to document the structural features, physical associations, and atmospheric metamorphosis of irregular snow crystals. The crystals appear as irregular hexagons, measuring 60 to 90 mm across, when viewed from the a-axis. Their length (c-axis) rarely exceeds the diameter. The irregular crystals are occasionally found as secondary particles on other larger forms of snow crystals; however, they most frequently occur in aggregates consisting of more than 100 irregular crystals. In the aggregates, the irregular crystals have their axes oriented parallel to one another and, collectively, tend to form columnar structures. Occasionally, these columnar structures exhibit rounded faces along one side, suggesting atmospheric metamorphoses during formation and descent. In extreme cases of metamorphoses, the aggregates would be difficult to distinguish from graupel. Frost, consisting of irregular crystals, has also been encountered, suggesting that atmospheric conditions that favor their growth can also occur terrestrially.