Non-native consumers can significantly alter processes at the population, community, and ecosystem level, and they are a major concern in many aquatic systems. Although the community-level effects of non-native anuran tadpoles are well understood, their ecosystem-level effects have been less studied. Here, I tested the hypothesis that natural densities of non-native bullfrog tadpoles (Lithobates catesbeianus) and native Woodhouse's toad tadpoles (Anaxyrus woodhousii) have dissimilar effects on aquatic ecosystem processes because of differences in grazing and nutrient recycling (excretion and egestion). I measured bullfrog and Woodhouse's carbon, nitrogen, and phosphorus nutrient recycling rates. Then, I determined the impact of tadpole grazing on periphyton biomass (chlorophyll a) during a 39-day mesocosm experiment. Using the same experiment, I also quantified the effect of tadpole grazing and nutrient excretion on periphyton net primary production (NPP). Lastly I measured how dissolved and particulate nutrient concentrations and respiration rates changed in the presence of the two tadpole species. Per unit biomass, I found that bullfrog and Woodhouse's tadpoles excreted nitrogen and phosphorus at similar rates, though Woodhouse's tadpoles egested more carbon, nitrogen, and phosphorus. However, bullfrogs recycled nutrients at higher N:C and N:P ratios. Tadpole excretion did not cause a detectable change in dissolved nutrient concentrations. However, the percent phosphorus in mesocosm detritus was significantly higher in both tadpole treatments, compared to a tadpole-free control. Neither tadpole species decreased periphyton biomass through grazing, although bullfrog nutrient excretion increased areal NPP. This result was due to higher biomass, not higher biomass-specific productivity. Woodhouse's tadpoles significantly decreased respiration in the mesocosm detritus, while bullfrog tadpoles had no effect. This research highlights functional differences between species by showing non-native bullfrog tadpoles and native Woodhouse's tadpoles may have different effects on arid, aquatic ecosystems. Specifically, it indicates bullfrog introductions may alter primary productivity and particulate nutrient dynamics.
First, a spatially collapsed model is used to develop voltage gated ion channels to study the excitability of the model neuron. Changing the channel densities reproduces different in situ observed firing patterns and induces a switch from resonator to integrator properties. Second, morphologically realistic multicompartment models are studied to investigate the passive properties of MN5. The passive electrical parameters fall in a range that is commonly observed in neurons, MN5 is spatially not compact, but for the single subtrees synaptic efficacy is location independent. Further, different subtrees are electrically independent from each other. Third, a continuum approach is used to formulate a new cable theoretic model to study the output in a dendritic cable with many subtrees, both analytically and computationally. The model is validated, by comparing it to a corresponding model with discrete branches. Further, the approach is demonstrated using MN5 and used to investigate spatially distributions of voltage gated ion channels.
Humans have dramatically increased phosphorus (P) availability in terrestrial and aquatic ecosystems. As P is often a limiting nutrient of primary production, changes in its availability can have dramatic effects on ecosystem processes. I examined the effects of calcium carbonate (CaCO3) deposition, which can lower P concentrations via coprecipitation of phosphate, on P availability in two systems: streams in the Huachuca Mountains, Arizona, and a stream, Río Mesquites, in Cuatro Ciénegas, México. Calcium carbonate forms as travertine in the former and within the microbialites of the latter. Despite these differences, CaCO3 deposition led to lowered P availability in both systems. By analyzing a three-year dataset of water chemistry from the Huachuca Mountain streams, I determined that P concentrations were negatively related to CaCO3 deposition rates. I also discovered that CaCO3 was positively correlated with nitrogen concentrations, suggesting that the stoichiometric effect of CaCO3 deposition on nutrient availability is due not only to coprecipitation of phosphate, but also to P-related constraints on biotic nitrogen uptake. Building from these observations, bioassays of nutrient limitation of periphyton growth suggest that P limitation is more prevalent in streams with active CaCO3 deposition than those without. Furthermore, when I experimentally reduced rates of CaCO3 deposition within one of the streams by partial light-exclusion, areal P uptake lengths decreased, periphyton P content and growth increased, and periphyton nutrient limitation by P decreased. In Río Mesquites, CaCO3 deposition was also associated with P limitation of microbial growth. There, I investigated the consequences of reductions in CaCO3 deposition with several methods. Calcium removal led to increased concentrations of P in the microbial biomass while light reductions decreased microbial biomass and chemical inhibition had no effect. These results suggest that CaCO3 deposition in microbialites does limit biological uptake of P, that photoautotrophs play an important role in nutrient acquisition, and, combined with other experimental observations, that sulfate reduction may support CaCO3 deposition in the microbialite communities of Río Mesquites. Overall, my results suggest that the effects of CaCO3 deposition on P availability are general and this process should be considered when managing nutrient flows across aquatic ecosystems.
A mathematical model is developed for the spread of rabies in a spatially distributed fox population to model the spread of the rabies epizootic through middle Europe that occurred in the second half of the 20th century. The model considers both territorial and wandering rabid foxes and includes a latent period for the infection. Since the model assumes these two kinds of rabid foxes, it is a system of both partial differential and integral equations (with integration
over space and, occasionally, also over time). To study the spreading speeds of the rabies epidemic, the model is reduced to a scalar Volterra-Hammerstein integral equation, and space-time Laplace transform of the integral equation is used to derive implicit formulas for the spreading speed. The spreading speeds are discussed and implicit formulas are given for latent periods of fixed length, exponentially distributed length, Gamma distributed length, and log-normally distributed length. A number of analytic and numerical results are shown pertaining to the spreading speeds.
Further, a numerical algorithm is described for the simulation
of the spread of rabies in a spatially distributed fox population on a bounded domain with Dirichlet boundary conditions. I propose the following methods for the numerical approximation of solutions. The partial differential and integral equations are discretized in the space variable by central differences of second order and by
the composite trapezoidal rule. Next, the ordinary or delay differential equations that are obtained this way are discretized in time by explicit
continuous Runge-Kutta methods of fourth order for ordinary and delay differential systems. My particular interest
is in how the partition of rabid foxes into
territorial and diffusing rabid foxes influences
the spreading speed, a question that can be answered by purely analytic means only for small basic reproduction numbers. I will restrict the numerical analysis
to latent periods of fixed length and to exponentially
distributed latent periods.
The results of the numerical calculations
are compared for latent periods
of fixed and exponentially distributed length
and for various proportions of territorial
and wandering rabid foxes.
The speeds of spread observed in the
simulations are compared
to spreading speeds obtained by numerically solving the analytic formulas
and to observed speeds of epizootic frontlines
in the European rabies outbreak 1940 to 1980.