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.
explained by natural selection acting on traits that persisted "for the good of the group" prompted a series of debates about group-level selection and the effectiveness with which natural selection could act at or across multiple levels of biological organization. For some this topic remains contentious, while others consider the debate settled, even while disagreeing about when and how resolution occurred, raising the question: "Why have these debates continued?"
Here I explore the biology, history, and philosophy of the possibility of natural selection operating at levels of biological organization other than the organism by focusing on debates about group-level selection that have occurred since the 1960s. In particular, I use experimental, historical, and synthetic methods to review how the debates have changed, and whether different uses of the same words and concepts can lead to different interpretations of the same experimental data.
I begin with the results of a group-selection experiment I conducted using the parasitoid wasp Nasonia, and discuss how the interpretation depends on how one conceives of and defines a "group." Then I review the history of the group selection controversy and argue that this history is best interpreted as multiple, interrelated debates rather than a single continuous debate. Furthermore, I show how the aspects of these debates that have changed the most are related to theoretical content and empirical data, while disputes related to methods remain largely unchanged. Synthesizing this material, I distinguish four different "approaches" to the study of multilevel selection based on the questions and methods used by researchers, and I use the results of the Nasonia experiment to discuss how each approach can lead to different interpretations of the same experimental data. I argue that this realization can help to explain why debates about group and multilevel selection have persisted for nearly sixty years. Finally, the conclusions of this dissertation apply beyond evolutionary biology by providing an illustration of how key concepts can change over time, and how failing to appreciate this fact can lead to ongoing controversy within a scientific field.
During 2013 and 2014, vegetation and herpetofauna were monitored at 21 riparian locations along the Virgin River via trapping and visual encounter surveys. Study sites were divided into four stand types based on density and percent cover of dominant trees (Tamarix, Prosopis, Populus, and Salix) and presence of restoration activities: Tam, Tam-Pros, Tam-Pop/Sal, and Restored Tam-Pop/Sal. Restoration activities consisted of mechanical removal of non-native trees, transplanting native trees, and introduction of water flow. All sites were affected by biological control. I predicted that herpetofauna abundance would vary between stand types and that herpetofauna abundance would be greatest in Restored Tam-Pop/Sal sites due to increased habitat openness and variation following restoration efforts.
Results from trapping indicated that Restored Tam-Pop/Sal sites had three times more total lizard and eight times more Sceloporus uniformis captures than other stand types. Anaxyrus woodhousii abundance was greatest in Tam-Pop/Sal and Restored Tam-Pop/Sal sites. Visual encounter surveys indicated that herpetofauna abundance was greatest in the Restored Tam-Pop/Sal site compared to the adjacent Unrestored Tam-Pop/Sal site. Habitat variables were reduced to six components using a principle component analysis and significant differences were detected among stand types. Restored Tam-Pop/Sal sites were most similar to Tam-Pop/Sal sites. S. uniformis were positively associated with large woody debris and high densities of Populus, Salix, and large diameter Prosopis.
Restored Tam-Pop/Sal sites likely supported higher abundances of herpetofauna, as these areas exhibited greater habitat heterogeneity. Restoration activities created a mosaic habitat by reducing canopy cover and increasing native tree density and surface water. Natural resource managers should consider implementing additional restoration efforts following biological control when attempting to restore riparian areas dominated by Tamarix and other non-native trees.