General methods for sensitivity analysis of equilibrium dynamics in patch occupancy models

Sensitivity analysis is a useful tool for the study of ecological models that has many potential applications for patch occupancy modeling. Drawing from the rich foundation of existing methods for Markov chain models, I demonstrate new methods for sensitivity analysis of the equilibrium state dynamics of occupancy models. Estimates from three previous studies are used to illustrate the utility of the sensitivity calculations: a joint occupancy model for a prey species, its predators, and habitat used by both; occurrence dynamics from a well-known metapopulation study of three butterfly species; and Golden Eagle occupancy and reproductive dynamics. I show how to deal efficiently with multistate models and how to calculate sensitivities involving derived state variables and lower-level parameters. In addition, I extend methods to incorporate environmental variation by allowing for spatial and temporal variability in transition probabilities. The approach used here is concise and general and can fully account for environmental variability in transition parameters. The methods can be used to improve inferences in occupancy studies by quantifying the effects of underlying parameters, aiding prediction of future system states, and identifying priorities for sampling effort.     

Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.     

No evidence for homeoviscous adaptation in intertidal snails: analysis of membrane fluidity during thermal acclimation, thermal acclimatization, and across thermal microhabitats

Many eurythermal organisms alter composition of their membranes to counter perturbing effects of environmental temperature variation on membrane fluidity, a process known as homeoviscous adaptation. Marine intertidal gastropods experience uniquely large thermal excursions that challenge the functional integrity of their membranes on tidal and seasonal timescales. This study measured and compared membrane fluidity in marine intertidal snail species under three scenarios: (1) laboratory thermal acclimation, (2) thermal acclimatization during a hot midday low tide, and (3) thermal acclimatization across the vertical intertidal zone gradient in temperature. For each scenario, we used fluorescence polarization of the membrane probe DPH to measure membrane fluidity in individual samples of gill and mantle tissue. A four-week thermal acclimation of Tegula funebralis to 5, 15, and 25°C did not induce differences in membrane fluidity. Littorina keenae sampled from two thermal microhabitats at the beginning and end of a hot midday low tide exhibited no significant differences in membrane fluidity, either as a function of time of day or as a function of thermal microhabitat, despite changes in body temperature up to 24°C within 8 h. Membrane fluidities of a diverse group of snails collected from high, middle, and low vertical regions of the intertidal zone varied among species but did not correlate with thermal microhabitat. Our data suggest intertidal gastropod snails do not exhibit homeoviscous adaptation of gill and mantle membranes. We discuss possible alternatives for how these organisms counter thermal excursions characteristic of the marine intertidal zone.     

Resource availability and the abundance of an N-based defense in Australian tropical rain forests

Plant defense theories predict that relatively resource-rich environments (those with more fertile soil) will support a greater abundance of plants with nitrogen-based chemical defense, but this has yet to be adequately tested. We tested this prediction by measuring the diversity and contribution to total biomass of cyanogenic plants (those that release hydrogen cyanide from endogenous cyanide-containing compounds) in the Australian tropical rain forest. We examined 401 species in thirty 200-m2 plots, six at each of five sites, for cyanogenesis. In upland/highland rain forest, two pairs of sites similar in rainfall and altitude, but differing in soil nutrients, were selected, as well as one site in lowland rain forest. Sites differed markedly in species composition and foliar N was positively related to soil fertility. Holding altitude constant, we did not detect significant differences in the proportion of cyanogenic species with soil fertility, nor did we consistently detect significant increases in the contribution of cyanogenic species to total biomass on higher nutrient sites. Thus we found no clear evidence that soil fertility affects the distribution and prevalence of species investing in a constitutive N-based defense at the community level.     

Temporal changes in C and N stocks of restored prairie: implications for C sequestration strategies

The recovery of ecosystem C and N dynamics after disturbance can be a slow process. Chronosequence approaches offer unique opportunities to use space-for-time substitution to quantify the recovery of ecosystem C and N stocks and estimate the potential of restoration practices for C sequestration. We studied the distribution of C and N stocks in two chronosequences that included long-term cultivated lands, 3- to 26-year-old prairie restorations, and remnant prairie on two related soil series. Results from the two chronosequences did not vary significantly and were combined. Based on modeling predictions, the recovery rates of different ecosystem components varied greatly. Overall, C stocks recovered faster than N stocks, but both C and N stocks recovered more rapidly for aboveground vegetation than for any other ecosystem component. Aboveground C and N reached 95% of remnant levels in only 13 years and 21 years, respectively, after planting to native vegetation. Belowground plant C and N recovered several decades later, while microbial biomass C, soil organic C (SOC), and total soil N recovered on a century timescale. In the cultivated fields, SOC concentrations were depleted within the surface 25 cm, coinciding with the depth of plowing, but cultivation apparently led to redistribution of soil C, increasing SOC stocks deeper in the soil profile. The restoration of prairie vegetation was effective at rebuilding soil organic matter (SOM) in the surface soil. Accrual rates were maintained at 43 g C x m(-2) x yr(-1) and 3 g N x m(-2) x yr(-1) in the surface 0.16 Mg/m2 soil mass during the first 26 years of restoration and were predicted to reach 50% of their storage potential (3500 g C/m2) in the first 100 years. We conclude that restoration of tallgrass prairie vegetation can restore SOM lost through cultivation and has the potential to sequester relatively large amounts of SOC over a sustained period of time. Whether restored prairies can retain the C apparently transferred to the subsoil by cultivation practices remains to be seen.     

Plant uptake of inorganic and organic nitrogen: neighbor identity matters

The importance of interspecific competition as a cause of resource partitioning among species has been widely assumed but rarely tested. Using neighbor removals in combination with 15N tracer additions in the field, we examined variation among three alpine species in the uptake of 15N-NH4+, 15N-NO3-, and 15N-13C-[2]-glycine in intact neighborhoods, when paired with a specific neighbor, and when all neighbors were removed. Species varied in the capacity to take up 15N-labeled NH4+, NO3-, and glycine in intact neighborhoods and in interspecific pairs. When interspecific neighbor pairs were compared with no neighbor controls, neighbors reduced 15N uptake in target species by as much as 50%, indicating competition for N. Furthermore, neighbor identity influenced the capacity of species to take up different forms of N. Thus, competition within interspecific neighbor pairs often caused reduced uptake of a particular form of N, as well as shifts to uptake of an alternative form of N. Such shifts in resource use as a result of competition are an implicit assumption in studies of resource partitioning but have rarely been documented. Our study suggests that plasticity in the uptake of different forms of N may be a mechanism by which cooccurring plants reduce competition for N.     

Hydrological properties of a clay loam soil after long-term cattle manure application

Limited information exists on the effect of long-term application of beef cattle (Bos taurus) manure on soil hydrological properties in the Great Plains region of North America. A site on a clay loam soil (Typic Haploboroll) was used to examine the effect of manure addition on selected soil hydrological properties in 1997 and 1998. The manure was annually applied in the fall for 24 yr at one, two, and three times the recommended rates (in 1973) under dryland (0, 30, 60, and 90 Mg ha(-1) wet basis) and irrigation (0, 60, 120, and 180 Mg ha(-1)). Manure significantly (P < or = 0.05) increased soil water retention (0-5 and 10-15 cm) by 5 to 48% compared with the control at most potentials between 0 and -1500 kPa. Field soil water content (0-5 and 10-15 cm) was increased by 10 to 22% in the summers of 1997 and 1998. Manure increased ponded infiltration by more than 200% at 90 Mg ha(-1) under dryland (1998) and at rates > or = 120 Mg ha(-1) under irrigation (1997). Field-saturated hydraulic conductivity (Kfs) of surface soil (1-cm depth) was significantly increased by 76 to 128% under dryland (1998) and irrigation (1997), as were number of pores > 1120 microm in diameter (37-128% increase). In contrast, manure rate had little or no effect on unsaturated hydraulic conductivity [K(psi)] values (-0.3, -0.5, -0.7, and -1.0 kPa) in 1997 and 1998. Overall, soil hydrological parameters generally had a neutral or positive response to 24 yr of annual manure addition.     

Prescribed burns and wildfires in Colorado: impacts of mitigation measures on indoor air particulate matter

Wildfires and prescribed burns are receiving increasing attention as sources of fine particulate matter (PM2.5). The goal of this research project was to understand the impact of mitigation strategies for residences impacted by scheduled prescribed burns and wildfires. Pairs of residences were solicited to have PM2.5 concentrations monitored inside and outside of their houses during four fires. The effect of using air cleaners on indoor PM2.5 was investigated, as well as the effect of keeping windows closed. Appropriately sized air cleaners were provided to one of each pair of residences; occupants of all of the residences were asked to keep windows shut and minimize opening of exterior doors. Additionally, residents were asked to record all of the activities that may be a source of particulate matter, such as cooking and cleaning. Measurements were made during one prescribed burn and three wildfires during the 2002 fire season. Outdoor 24-hr average PM2.5 concentrations ranging from 6 to 38 microg/m3 were measured during the fires, compared with levels of 2-5 microg/m3 during background measurements when no fires were burning. During the fires, PM2.5 was < 3 microg/m3 inside all of the houses with air cleaners installed. This corresponds with a decrease of 63-88% in homes with the air cleaners operating when compared with homes without air cleaners. In the homes without the air cleaners, measured indoor concentrations were 58-100% of the concentrations measured outdoors.     

Nutrient input and removal trends for agricultural soils in nine geographic regions in Arkansas

Knowledge of the balance between nutrient inputs and removals is required for identifying regions that possess an excess or deficit of nutrients. This assessment describes the balance between the agricultural nutrient inputs and removals for nine geographical districts within Arkansas from 1997 to 2001. The total N, P, and K inputs were summed for each district and included inorganic fertilizer and collectable nutrients excreted as poultry, turkey, dairy, and hog manures. Nutrients removed by harvested crops were summed and subtracted from total nutrient inputs to calculate the net nutrient balance. The net balances for N, P, and K were distributed across the hectarage used for row crop, hay, pasture, or combinations of these land uses. Row-crop agriculture predominates in the eastern one-third and animal agriculture predominates in the western two-thirds of Arkansas. Nutrients derived from poultry litter accounted for >92% of the total transportable manure N, P, and K. The three districts in the eastern one-third of Arkansas contained 95% of the row-crop hectarage and had net N and P balances that were near zero or negative. The six districts in the western two-thirds of Arkansas accounted for 89 to 100% of the animal populations, had positive net balances for N and P, and excess P ranged from 1 to 9 kg P ha(-1) when distributed across row-crop, hay, and pasture hectarage. Transport of excess nutrients, primarily in poultry litter, outside of the districts in western Arkansas is needed to achieve a balance between soil inputs and removals of P and N.     

Bermudagrass sod growth and metal uptake in coal combustion by-product-amended media

Coal combustion by-products (CCB) include fly ash and bottom ash and are generated nationally at rates of 10(8) Mg yr(-1). Land applications of CCB have improved physicochemical properties of soil, yet inherent bulkiness and trace metal content of CCB often limit their use. Likewise, utilization of biosolids and manure as fertilizer can be problematic due to unfavorable nutrient ratios. A 2-yr field study evaluated environmental and technical parameters associated with CCB-organic waste utilization as growth media in turfgrass sod production. Experimental growth media formulated with CCB and organic waste and a sand-compost control mixture were uniformly spread at rates from 200 to 400 m3 ha(-1) and sprigged with hybrid bermudagrass [Cynodon dactylon (L.) Pers. x C. transvaalensis Burtt-Davy]. Leaf clippings were collected and analyzed for total elemental content each year. In Year 2, growth media samples were collected during establishment 47 and 84 days after planting (DAP) and viable Escherichia coli organisms were quantified. At harvest (99 or 114 DAP), sod biomass and physicochemical properties of the growth media were measured. During sod propagation, micronutrient and metal content in leaf clippings varied by growth media and time. After 47 d of typical sod field management, viable E. coli pathogens were detected in only one biosolids-amended plot. No viable E. coli were measured at 84 DAP. In both years, sod biomass was greatest in media containing biosolids and fly ash. Following installation of sod, evaluations did not reveal differences by media type or application volume. Using CCB-organic waste mixes at the rates described herein is a rapid and environmentally safe method of bermudagrass sod production.