Data set ID:
Introduction. Animal consumers have important roles in ecosystems (Chew 1974, 1976), determining plant species composition and structure (Harper 1969, Pacala and Crawley 1992, Crawley 1983, 1989), regulating rates of plant production and nutrient cycling (Naiman 1988, McNaughton et al. 1989, Holland et al. 1992), and altering soil structure and chemistry (Milchunas et al. 1993, Huntly 1991). Desertification of semi-arid grasslands in the Southwest United States by domestic livestock provides an important example of herbivore regulation of ecosystem structure and function (Schlesinger et al. 1990). The species composition and physical structure of these desert grassland ecosystems were significantly altered by alien herbivores about 100 years ago (Bahre 1991, York and Dick-Peddie 1968, Gardner 1951, Hastings and Turner 1980, Buffington and Herbel 1965, Dick-Peddie 1993). To what extent the spatial patterns of semi-arid shrubland and grassland plant production and soil characteristics are currently controlled by plant resource use, abiotic factors, or consumers is not known. Desertification is an ecosystem-level phenomenon occurring on a global scale with great relevance to human welfare (Nelson 1988). In order to understand the processes that contribute to desertification, we must fully understand interactions among the components of arid-land ecosystems. Schlesinger et al. (1990) suggest that in the absence of continued livestock perturbations, plant resource use and abiotic factors appear to be the principal factors accounting for the persistence of desert shrublands in desertified semi-arid grasslands. However, Brown and Heske (1990a) provide evidence that indigenous small mammal consumers may also have a major role in determining vegetation structure in those desert ecosystems. Brown and Heske (1990a, Heske et al. 1993) found that the exclusion of rodents from Chihuahuan Desert creosotebush shrubland areas resulted in a significant increase in grass cover over a 12 year period. Brown and Heske (1990a) concluded that rodents were keystone species in those desert shrub communities, greatly influencing vegetation structure. Rodents are also known to have significant influences on plant species composition and diversity in desert communities (Inouye et al. 1980, Heske et al. 1993, Brown et al. 1986). Several species of granivorous rodents (Family: Heteromidae, genera: Dipodomys, Perognathus, Chaetodipus) appear to have the greatest influence on vegetation herbivory. Soil disturbance through the digging activities of rodents can have profound local effects on plant species composition and vegetation structure in the Chihuahuan Desert (Moroka et al. 1982). Digging activities of desert rodents intermix surface soils with subsurface soils (Abaturov 1972), and increase rainfall infiltration (Soholt 1975). Reported measures of the percentage of desert soil surface areas disturbed by rodent digging activities in desert enviroments range from 10% (Abaturov 1972) to 4.5% (Soholt 1975). Burrowing activities increase local soil nutrient and water status, creating favorable sites for increased plant densities, biomass production, and increased species diversity (Morehead et al. 1989, Mun and Whitford 1990). Rabbits (Lagomorpha: Black-tailed jackrabbits, Lepus californicus, and desert cottontail rabbits, Sylvilagus aduboni) are also important consumers of desert vegetation (Brown 1947, Johnson & Anderson 1984, Steinberger and Whitford 1983, Ernest 1994). Rabbits can have significant effects on plant species composition and structure resulting from selective herbivory (Gibbens et al. 1993, Clark and Wagner 1984, Norris 1950, Zeevalking and Fresco 1977). Gibbens et al. (1993) found that excluding rabbits from Chihuahuan Desert creosotebush (Larrea tridentata) communities over a period of 50 years increased the canopy cover of some grasses, and also increased canopy cover of some shrub species. Small mammal (rodent and rabbit) populations may fluctuate considerably with variation in climate and annual plant production (Brown et al. 1979, Brown & Heske 1990, Brown & Zeng 1989, Whitford 1976, Johnson & Anderson 1984). Reproduction in desert rodents is known to be induced by plant foliage production (Reichman and Van De Graff 1975, Beatley 1969). If small mammals are keystone species affecting plant species composition and structure in desert ecosystems, then the impacts of small mammals on vegetation are probably linked with variation in climate and plant production. A reciprocal plant-herbivore/granivore feedback system may result, where small mammal populations and thus impacts on vegetation, are initially determined by climate influences on plant food resource availability to the small mammals. Thus, the effects of small mammals during dry years will probably be different from the effects during wet years because of different population sizes. If this is so, one should be able to measure differential effects of small mammals on plant communities over series of wet or dry years, such as El Nino and La Nina cycles (Nicholls 1988). Such reciprocal interactions should also occur in relation to long-term (decades) climate change. The effects of any one small mammal species population on the biotic community will be complicated by competitive interactions with other mammal species (Munger & Brown 1981, Brown & Zeng 1989, Brown & Heske 1990), however overall impacts on vegetation and soils by the combined effects of all small mammal species may be closely linked with variation in precipitation and plant production. Depending upon the persistence of plant food resources such as foliage or seeds, lag times in consumer impacts may be expected following periods of precipitation and plant production. In desert ecosystems, widely scattered shrubs produce a patch pattern of fertile islands with high plant biomass production and soil nutrients, surrounded by relatively unproductive barren soil (West and Klemmedson 1978, Crawford and Gosz 1982). Researchers at the Jornada Long-Term Ecological Research site in New Mexico have proposed a desertification model suggesting that perturbations caused by domestic livestock grazing and climate change initiated processes transforming grasslands with relatively homogeneous resource distributions to shrubland environments with relatively heterogenous resource distributions (Schlesinger et al. 1990). This patchy vegetation/resource distribution pattern is stable under present climate regimes, and appears to be maintained by plant resource use and abiotic soil processes (Schlesinger et al. 1990). However, Wagner (1976, page 195) suggested that small mammals were probably maintaining shrubland dominated ecosystems at the Jornada by suppressing grasses through selective herbivory. Research Hypotheses. The purpose of this study is to determine whether or not the activities of small mammals regulate plant community structure, plant species diversity, and spatial vegetation patterns in Chihuahuan Desert shrublands and grasslands. What role if any do indigenous small mammal consumers have in maintaining desertified landscapes in the Chihuahuan Desert? Additionally, how do the effects of small mammals interact with changing climate to affect vegetation patterns over time? This study will provide long-term experimental tests of the roles of consumers on ecosystem pattern and process across a latitudinal climate gradient. The following questions or hypotheses will be addressed. 1) Do small mammals influence patterns of plant species composition and diversity, vegetation structure, and spatial patterns of vegetation canopy cover and biomass in Chihuahuan Desert shrublands and grasslands? Are small mammals keystone species that determine plant species composition and physiognomy of Chihuahuan Desert communities as Brown and Heske (1990a) and Gibbens et al. (1993) suggest? Do small mammals have a significant role in maintaining the existence of shrub islands and spatial heterogeneity of creosotebush shrub communities? 2) Do small mammals affect the taxonomic composition and spatial pattern of vegetation similarly or differently in grassland communities as compared to shrub communities? How do patterns compare between grassland and shrubland sites, and how do these relatively small scale patterns relate to overall landscape vegetation patterns? 3) Do small mammals interact with short-term (annual) and long-term (decades) climate change to affect temporal changes in vegetation spatial patterns and species composition? Other Consumers. Ants are important consumers in Chihuahuan Desert ecosystems (MacKay 1991), and granivorous ants are known to have competitive interactions with rodents (Brown & Davidson 1977, Brown et al. 1979) for plant seed resources. Termites are important detritivores in Chihuahuan Desert ecosystems (MacKay 1991) and appear to have key roles in plant litter decomposition and nutrient cycling (Whitford et al. 1982, Schaefer & Whitford 1981), and in altering soil structure and hydrologic processes (Elkins et al. 1986). Grasshoppers are important herbivores in Chihuahuan Desert ecosystems (Rivera 1986, Wisdom 1991, Richman et al. 1993), with various species specializing on most of the different plant species present in any location (Otte 1976, Joern 1979). Since manipulations of small mammals will probably affect these arthropod consumers, we will monitor these other consumers on the measurement plots to document any changes. Documentation of changes or lack of changes in ant, termite, and grasshopper consumer groups will be needed to interpret the results of small mammal manipulations on vegetation and soils. For example, if removal of rodents results in an increase of seed-harvesting ants, changes or lack of changes in vegetation and soils may be attributed to compensatory granivory from the increase in ants. Small mammals are the consumer group that appears to have the greatest influence on Chihuahuan Desert communities (see literature citations above). Given the known ecological importance of small mammals and the complexity and difficulties that would be associated with manipulating small mammals and arthropods, we have chosen to start with experiments on small mammals first. If these other consumer groups appear to have important interactions with small mammals, we will pursue additional experiments in the future to focus on those interactions, and to elucidate the ecological roles of these arthropod consumers.
Field micro-cassette tape recorders
Study Site and Experimental Design Description A creosotebush shrub study site and a black grama grassland study site have been established at each of the Sevilleta, Jornada and Mapimi research locations. Each study site is 1 km by 0.5 km in area. Three rodent trapping webs and four replicate experimental blocks of plots are randomly located at each study site to measure vegetation responses to the exclusion of small mammals (see Figures 1 & 2 for diagrams of the two Jornada sites). The blocks of study plots are all oriented on a site in a X/Y coordinate system, with the access road to each site forming the X axis. The compass orientation at the Jornada grassland site is to the north, and Jornada creosotebush site is oriented to the south. Treatments within each block include one unfenced control plot (Treatment: C; control), one plot fenced with hardware cloth and poultry wire to exclude rodents and rabbits (Treatment: R; rodent), and one plot fenced only with poultry wire to exclude rabbits (Treatment: L; lagomorph), and one plot fenced with barbed wire to exclude cattle (Treatment B; bovine). Note that there are cattle exclosure plots only at the Jornada grassland site where cattle are present, for a total of 4 measurement plots at each of the grassland site blocks. There are only 3 measurement plots at each of the creosotebush site blocks. The treatments were randomly assigned to each of the four possible plots in each block independently, and their arrangements differ from block to block. Each of the plots in a replicate block are separated by 20 meters (see Figure 3). Each experimental measurement plot measures 36 meters by 36 meters (see Figure 4). A grid of 36 sampling points are positioned at 5.8-meter intervals on a systematically located 6 by 6 point grid within each plot. A permanent one-meter by one-meter vegetation measurement quadrat is located at each of the 36 points. The 36 quadrats are numbered 1-36, starting with number 1 in the top left corner of each plot, and running left to right, then down one row, and then right to left, and so on (see Figure 4). A 2-foot rebar marks the lower right corner of each quadrat, and an aluminum tag on the rebar gives the quad number. 3-inch nails were originally placed in the top left corner of each quadrat. These may be difficult to see. A 3-meter wide buffer area is situated between the grid of 36 points and the perimeter of each plot. Working on the Study Plots Always avoid walking on the quadrats, and do not walk across plots or anywhere within a block unless absolutely necessary. We are attempting to measure the effects of rodents and rabbits on plants and soils, not the effects of humans. When working on a plot, always walk on a line just below the rebar and quadrats, or on a line to the right of the rebar and quadrats (see dotted lines in Figure 4). Please try to walk gently and flat-footed on the study plots to minimize soil disturbance. When placing the vegetation measurement frame on a quadrat, be careful not to disturb the soil with the frame legs or your feet. When leaning over the measurement frame, be careful not to put your foot on the quadrat. We must measure all human caused disturbances to the soil surface of each quadrat. Vegetation Quadrat Measurements The foliage canopy area and maximum height of each plant species is measured from each quadrat. Several other variables are also measured from each quadrat, including soil surface disturbance, soil surface leaf litter cover, soil surface cryptogam crust cover, termite mud casing, and number of rabbit feces. All cover values are measured from the vegetation measurement frame, which is 1 meter by 1 meter, and partitioned into a grid of 100, 10 cm by 10 cm squares. Cover is measured by counting the number of 10cm squares that are occupied by the foliage canopy of a particular plant species, or by the soil disturbance, leaf litter etc. Portions of the 10 cm squares are also measured, down to 0.1 of a square. Detailed descriptions of measurement techniques are given below for each of the different variables measured. Placement of the vegetation quadrat measurement frame Walk from quadrat to quadrat along the lines mentioned above and marked in Figure 4. When you reach a quadrat, place one leg of the frame immediately next to and touching the 2-foot rebar, over the quadrat. Place the leg for the opposite corner of the frame (from bottom to top of a diamond shape) just inside of the 3-inch nail located at a 315 degree angle from the rebar on the plot X/Y, and compass coordinate system. The frame should be positioned so that the sides are parallel to the sides of the plot and directly over the quadrat, and in line with the rows of rebar (see Figure 4). You may not be able to see the nail, if not, line the sides of the frame with the rows of rebar the best you can. Recording Data You will record the quadrat data into your tape recorder. Each time you start recording data from a new plot (even if you are just helping someone else finish a plot), always start your recording by stating: "These are vegetation quadrat measurements for the Jornada small mammal exclosure study." Then state the date (month, day, year), the site (G or C), the block (1,2, 3, or 4), the plot (1, 2, 3, or 4), and the treatment (C, R, or L). If you are helping someone else on a plot, state so, and record who the other person is. Go to the first quadrat you will be taking measurements from, and record the quadrat number (1-36) before you start recording data, by stating "starting quadrat (n)". Note that (n) means whatever the appropriate number or letter is. When you are finished recording data for that quadrat, state "finished with quadrat (n)." Then go on to the next quadrat, and so on. When you have finished collecting data from the plot, state "finished with quadrat measurements on plot (n), of block (n), at site (n), on (date)." Additional Observations, Corrections, and Comments If you have already recorded data for a variable, e.g., rabbit feces, but while looking over the quadrat later, see some more, you may either make a correction to the previously recorded values, or make another observation (e.g., "correction, rabbit feces count (n)", or "rabbit feces count (n)." At any time, if you realize that you recorded incorrect data, just state "correction, ....." Always start corrections with the statement "correction". Do not try to run-back the tape and re-record over mistakes, just state "correction", and mention the problem. For comments, state "comment" and then record whatever you want to say about something. Comments are useful if you are uncertain about something, or see something unusual. Try to minimize use of comments though. Checking Your Tape Recordings Be sure to frequently play-back a small portion of your tape to verify that the tape recorder is working properly. Stop and play-back at least at the end of each line of 6 quadrats. Checking your tape after every other quadrat is even better. Labeling Your Tapes Be sure to label your tape, both on the cardboard case insert, and directly on the tape. Use the following format to label your tape: e.g., SMESVQF96-DL2 Where: SMES = small mammal exclosure study VQ = vegetation quadrat data F96 = fall 1996, or S96 for spring 1996 DL2 = initials for your name, in this case Dave Lightfoot, and the number of the ape, in this case, my second tape. Procedures for Vegetation Quadrat Measurements Below is a listing of procedures and values and ranges of values that you should record for each of the variables measured on the vegetation quadrats. IMPORTANT NOTE!!! Be sure to record an entry for all six of the following variables, even if the measurement or count is zero. So, for example, if there are no rabbit feces, record "rabbit feces count zero" or if there is no termite casing, record "termite casing zero" etc. Get into a routine of examining each quadrat for each of the 6 variables. For example, start with plant cover, starting with the dominant plant species, then look for leaf litter, then look for cryptogams, then look for soil disturbance, then look for rabbit feces, then look for termite casing. And be sure to record a value for each of the 6 variables for each quadrat. Leaf Litter Cover State "leaf litter cover(n)" for the cover of leaf litter on the soil surface of the quadrat in terms of the 10 cm squares. For cover values less than 5, use increments of 1.0. For cover values greater than 5, use increments of 5.0. Leaf litter includes all detached dead plant material on the soil surface, including woody branches. Only measure leaf litter cover that is in the open, do not attempt to measure within clumps of grass, etc. Some leaf litter cover has distinctive margins and is easy to define and measure. However, much leaf litter consists of many diffuse small patches that are separated by bare soil, and distributed throughout the quadrat. For such diffuse cover, determine the actual cover in one typical 10 by 10 cm square (e.g., 0.3), then count the number of squares with diffuse cover (e.g., 5), and multiply the number of squares by the actual cover for a typical square (e.g., 0.3 X 5 = 1.5, then round to 1.0 or 2.0, or if the value had been greater than 5, round to the nearest increment of 5.0 ) for the total litter cover. All litter cover is pooled into one observation, and no height is measured.
SAS programs will be used to analyze data
A creosotebush shrub study site and a black grama grassland study site have been established at each of the Sevilleta, Jornada and Mapimi research locations. Each study site is 1 km by 0.5 km in area. Three rodent trapping webs and four replicate experimental blocks of plots are randomly located at each study site to measure vegetation responses to the exclusion of small mammals. The blocks of study plots are all oriented on a site in a X/Y coordinate system, with the access road to each site forming the X axis. The compass orientation at the Jornada grassland site is to the north, and Jornada creosotebush site is oriented to the south. Treatments within each block include one unfenced control plot (Treatment: C; control), one plot fenced with hardware cloth and poultry wire to exclude rodents and rabbits (Treatment: R; rodent), and one plot fenced only with poultry wire to exclude rabbits (Treatment: L; lagomorph), and one plot fenced with barbed wire to exclude cattle (Treatment B; bovine). Note that there are cattle exclosure plots only at the Jornada grassland site where cattle are present, for a total of 4 measurement plots at each of the grassland site blocks. There are only 3 measurement plots at each of the creosotebush site blocks. The treatments were randomly assigned to each of the four possible plots in each block independently, and their arrangements differ from block to block. Each of the plots in a replicate block are separated by 20 meters. Each experimental measurement plot measures 36 meters by 36 meters (see Figure 4). A grid of 36 sampling points are positioned at 5.8-meter intervals on a systematically located 6 by 6 point grid within each plot. A permanent one-meter by one-meter vegetation measurement quadrat is located at each of the 36 points. The 36 quadrats are numbered 1-36, starting with number 1 in the top left corner of each plot, and running left to right, then down one row, and then right to left, and so on. A 2-foot rebar marks the lower right corner of each quadrat, and an aluminum tag on the rebar gives the quad number. 3-inch nails were originally placed in the top left corner of each quadrat. These may be difficult to see. A 3-meter wide buffer area is situated between the grid of 36 points and the perimeter of each plot. Rodent trapping webs are being used to determine the composition of rodent species at each study site, and to estimate densities of each species over time. The use of webs and distance measures to estimate rodent densities is statistically more robust than grid plot sampling and mark-release indices. Each rodent trapping web consists of a series of 12 equally spaced lines radiating from a central point. Each line consists of 12 trap stations. The first trap station is located 5 meters from the center, the next three at 5 meter intervals, and the remaining 8 at ten meter intervals. Each trap line is 100 meters long, and each web is 200 meters in diameter.
Twice each year through 1995: April and October Once every 5 years after 1995: October