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Description
During the 1960s, the long-standing idea that traits or behaviors could be
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.
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.
ContributorsDimond, Christopher C (Author) / Collins, James P. (Thesis advisor) / Gadau, Juergen (Committee member) / Laubichler, Manfred (Committee member) / Armendt, Brad (Committee member) / Lynch, John (Committee member) / Arizona State University (Publisher)
Created2014
Description
Infectious diseases have emerged as a significant threat to wildlife. Environmental change is often implicated as an underlying factor driving this emergence. With this recent rise in disease emergence and the acceleration of environmental change, it is important to identify the environmental factors that alter host-pathogen dynamics and their underlying mechanisms. The emerging pathogen Batrachochytrium dendrobatidis (Bd) is a clear example of the negative effects infectious diseases can have on wildlife. Bd is linked to global declines in amphibian diversity and abundance. However, there is considerable variation in population-level responses to Bd, with some hosts experiencing marked declines while others persist. Environmental factors may play a role in this variation. This research used populations of pond-breeding chorus frogs (Pseudacris maculata) in Arizona to test if three rapidly changing environmental factors nitrogen (N), phosphorus (P), and temperature influence the presence, prevalence, and severity of Bd infections. I evaluated the reliability of a new technique for detecting Bd in water samples and combined this technique with animal sampling to monitor Bd in wild chorus frogs. Monitoring from 20 frog populations found high Bd presence and prevalence during breeding. A laboratory experiment found 85% adult mortality as a result of Bd infection; however, estimated chorus frog densities in wild populations increased significantly over two years of sampling despite high Bd prevalence. Presence, prevalence, and severity of Bd infections were not correlated with aqueous concentrations of N or P. There was, however, support for an annual temperature-induced reduction in Bd prevalence in newly metamorphosed larvae. A simple mathematical model suggests that this annual temperature-induced reduction of Bd infections in larvae in combination with rapid host maturation may help chorus frog populations persist despite high adult mortality. These results demonstrate that Bd can persist across a wide range of environmental conditions, providing little support for the influence of N and P on Bd dynamics, and show that water temperature may play an important role in altering Bd dynamics, enabling chorus frogs to persist with this pathogen. These findings demonstrate the importance of environmental context and host life history for the outcome of host-pathogen interactions.
ContributorsHyman, Oliver J. (Author) / Collins, James P. (Thesis advisor) / Davidson, Elizabeth W. (Committee member) / Anderies, John M. (Committee member) / Elser, James J. (Committee member) / Escalante, Ananias (Committee member) / Arizona State University (Publisher)
Created2012
Description
Climate and land use change are projected to threaten biodiversity over the coming century. However, the combined effects of these threats on biodiversity and the capacity of current conservation networks to protect species' habitat are not well understood. The goals of this study were to evaluate the effect of climate change and urban development on vegetation distribution in a Mediterranean-type ecosystem; to identify the primary source of uncertainty in suitable habitat predictions; and to evaluate how well conservation areas protect future habitat in the Southwest ecoregion of the California Floristic Province. I used a consensus-based modeling approach combining three different species distribution models to predict current and future suitable habitat for 19 plant species representing different plant functional types (PFT) defined by fire-response (obligate seeders, resprouting shrubs), and life forms (herbs, subshurbs). I also examined the response of species grouped by range sizes (large, small). I used two climate models, two emission scenarios, two thresholds, and high-resolution (90m resolution) environmental data to create a range of potential scenarios. I evaluated the effectiveness of an existing conservation network to protect suitable habitat for rare species in light of climate and land use change. The results indicate that the area of suitable habitat for each species varied depending on the climate model, emission scenario, and threshold combination. The suitable habitat for up to four species could disappear from the ecoregion, while suitable habitat for up to 15 other species could decrease under climate change conditions. The centroid of the species' suitable environmental conditions could shift up to 440 km. Large net gains in suitable habitat were predicted for a few species. The suitable habitat area for herbs has a small response to climate change, while obligate seeders could be the most affected PFT. The results indicate that the other two PFTs gain a considerable amount of suitable habitat area. Several rare species could lose suitable habitat area inside designated conservation areas while gaining suitable habitat area outside. Climate change is predicted to be more important than urban development as a driver of habitat loss for vegetation in this region in the coming century. These results indicate that regional analyses of this type are useful and necessary to understand the dynamics of drivers of change at the regional scale and to inform decision making at this scale.
ContributorsBeltrán Villarreal, Bray de Jesús (Author) / Franklin, Janet (Thesis advisor) / Fenichel, Eli P (Committee member) / Kinzig, Ann P (Committee member) / Collins, James P. (Committee member) / Arizona State University (Publisher)
Created2012
Description
Novel resource inputs represent an increasingly common phenomenon in ecological systems as global change alters environmental factors and species distributions. In semiarid riparian areas, hydric pioneer tree species are being replaced by drought-tolerant species as water availability decreases. Additionally, introduced omnivorous crayfish, which feed upon primary producers, allochthonous detritus, and benthic invertebrates, can impact communities at multiple levels through both direct and indirect effects. In arid and semiarid systems of the American Southwest, crayfish may be especially important as detrital processors due to the lack of specialized detritivores. I tested the impact of virile crayfish (Orconectes virilis) on benthic invertebrates and detrital resources across a gradient of riparian vegetation drought-tolerance using field cages with leaf litter bags in the San Pedro River in Southeastern Arizona. Virile crayfish increased breakdown rate of drought-tolerant saltcedar (Tamarix ramosissima), but did not impact breakdown of Fremont cottonwood (Populus fremontii), Gooding's willow (Salix goodingii), or seepwillow (Baccharis salicifolia). The density and composition of the invertebrate community colonizing leaf litter bags were both heavily influenced by litter species but not directly by crayfish presence. As drought-tolerant species become more abundant in riparian zones, their litter will become a larger component of the organic matter budget of desert streams. By increasing breakdown rates of saltcedar, crayfish shift the composition of leaf litter in streams, which in turn may affect the composition and biomass of colonizing invertebrate communities. More research is needed to determine the full extent to which these alterations change community composition over time.
ContributorsMoody, Eric Kellan (Author) / Sabo, John L (Thesis advisor) / Collins, James P. (Committee member) / Stromberg, Juliet C. (Committee member) / Arizona State University (Publisher)
Created2012
Description
Primary production in aquatic ecosystems is often limited by the availability of nitrogen (N) and/or phosphorus (P). Animals can substantially alter the relative availability of these nutrients by storing and recycling them in differential ratios. Variation in these stoichiometric traits, i.e., the elemental phenotype, within a species can link organismal evolution to ecosystem function. I examined the drivers of intraspecific variation in the elemental phenotype of aquatic consumers to test for the generality of these effects. Over a thermal gradient in Panamá, I found that average specific growth grate and body P content of the mayfly Thraulodes increased with environmental temperature, but that these patterns were due to site-specific differences rather than the direct effects of warmer temperature. In a meta-analysis of published studies, I found that in fishes intraspecific variation in dietary N:P ratio had a significant effect on excretion N:P ratio, but only when accounting for consumption. I tested for the effects of variation in consumption on excretion N:P ratio among populations of the fish Gambusia marshi in the Cuatro Ciénegas basin in Coahuila, Mexico. G. marshi inhabits warm groundwater-fed springs where it often co-occurs with predatory fishes and cool runoff-dominated wetlands which lack predators. Using stoichiometric models, I generated predictions for how variation in environmental temperature and predation pressure would affect the N:P ratio recycled by fishes. Adult female G. marshi excretion N:P ratio was higher in runoff-dominated sites, which was consistent with predators driving increased consumption rates by G. marshi. This result was supported by a diet ration manipulation experiment in which G. marshi raised on an ad libitum diet excreted N:P at a lower ratio than fish raised on a restricted diet ration. To further support the impacts of predation on phenotypic diversification in G. marshi, I examined how body morphology varied among habitats and among closely related species. Both among and within species, predation had stronger effects on morphology than the physical environment. Overall, these results suggest that predation, not temperature, has strong effects on these phenotypic traits of aquatic consumers which can alter their role in ecosystem nutrient cycling through variation in consumption rates.
ContributorsMoody, Eric Kellan (Author) / Elser, James J (Thesis advisor) / Sabo, John L (Thesis advisor) / Angilletta, Michael J (Committee member) / Capps, Krista A (Committee member) / Collins, James P. (Committee member) / Arizona State University (Publisher)
Created2017
Description
Transgenic experiments in Drosophila have proven to be a useful tool aiding in the
determination of mammalian protein function. A CNS specific protein, dCORL is a
member of the Sno/Ski family. Sno acts as a switch between Dpp/dActivin signaling.
dCORL is involved in Dpp and dActivin signaling, but the two homologous mCORL
protein functions are unknown. Conducting transgenic experiments in the adult wings,
and third instar larval brains using mCORL1, mCORL2 and dCORL are used to provide
insight into the function of these proteins. These experiments show mCORL1 has a
different function from mCORL2 and dCORL when expressed in Drosophila. mCORL2
and dCORL have functional similarities that are likely conserved. Six amino acid
substitutions between mCORL1 and mCORL2/dCORL may be the reason for the
functional difference. The evolutionary implications of this research suggest the
conservation of a switch between Dpp/dActivin signaling that predates the divergence of
arthropods and vertebrates.
determination of mammalian protein function. A CNS specific protein, dCORL is a
member of the Sno/Ski family. Sno acts as a switch between Dpp/dActivin signaling.
dCORL is involved in Dpp and dActivin signaling, but the two homologous mCORL
protein functions are unknown. Conducting transgenic experiments in the adult wings,
and third instar larval brains using mCORL1, mCORL2 and dCORL are used to provide
insight into the function of these proteins. These experiments show mCORL1 has a
different function from mCORL2 and dCORL when expressed in Drosophila. mCORL2
and dCORL have functional similarities that are likely conserved. Six amino acid
substitutions between mCORL1 and mCORL2/dCORL may be the reason for the
functional difference. The evolutionary implications of this research suggest the
conservation of a switch between Dpp/dActivin signaling that predates the divergence of
arthropods and vertebrates.
ContributorsStinchfield, Michael J (Author) / Newfeld, Stuart J (Thesis advisor) / Capco, David (Committee member) / Laubichler, Manfred (Committee member) / Arizona State University (Publisher)
Created2019
Description
Emerging infectious diseases (EIDs) in vulnerable populations are a proposed cause of reduced global biodiversity due to local and regional extinctions. Chytridiomycosis, a fungal disease caused by Batrachochytrium dendrobatidis (Bd), is affecting amphibian populations worldwide.
Chapter 1 of this thesis reports using lab-raised larval tiger salamanders (Ambystoma tigrinum nebulosum), collected as eggs, to test if Bd infects them. Bd infects metamorphosed tiger salamanders; however, it is currently unknown if larvae can be infected by Bd. Adult frogs tend to host Bd on ventral surfaces and hind legs while tadpoles host Bd in keratinized mouthparts. No research has considered differences in infection between life stages of salamanders. It was hypothesized that Bd can colonize larvae in the same manner as metamorphosed animals. Larval salamanders were inoculated to test if Bd concentrations differ among body regions in larvae compared to metamorphosed salamanders. Larvae can carry Bd with the concentration of Bd varying between body region.
Chapter 2 report using native tiger salamanders (Ambystoma tigrinum nebulosum), from northern Arizona and Bd as a study system to test if Bd is native or introduced to Arizona. It was hypothesized that Bd is not endemic to Arizona, but is introduced. There are multiple hypotheses regarding potential routes Bd may have traveled through Arizona and into Mexico. These hypotheses were tested using the Kaibab Plateau in Coconino County, Arizona, as a study site. The plateau is isolated from surrounding areas by the Grand Canyon to the south and the Vermillion Cliffs to the north serving as major biogeographical barriers. It is hypothesized that tiger salamanders are not dispersing into or out of the Kaibab Plateau due to geological restrictions. Bd, therefore, should not be present on salamanders on the Kaibab Plateau due to geological restriction. Tiger salamanders in stock tanks located on the Kaibab as well as preserved museum specimens housed in the Arizona State University Natural History Collection were sampled. The results indicate that Bd occurs at low levels on Kaibab Plateau tiger salamanders.
Chapter 1 of this thesis reports using lab-raised larval tiger salamanders (Ambystoma tigrinum nebulosum), collected as eggs, to test if Bd infects them. Bd infects metamorphosed tiger salamanders; however, it is currently unknown if larvae can be infected by Bd. Adult frogs tend to host Bd on ventral surfaces and hind legs while tadpoles host Bd in keratinized mouthparts. No research has considered differences in infection between life stages of salamanders. It was hypothesized that Bd can colonize larvae in the same manner as metamorphosed animals. Larval salamanders were inoculated to test if Bd concentrations differ among body regions in larvae compared to metamorphosed salamanders. Larvae can carry Bd with the concentration of Bd varying between body region.
Chapter 2 report using native tiger salamanders (Ambystoma tigrinum nebulosum), from northern Arizona and Bd as a study system to test if Bd is native or introduced to Arizona. It was hypothesized that Bd is not endemic to Arizona, but is introduced. There are multiple hypotheses regarding potential routes Bd may have traveled through Arizona and into Mexico. These hypotheses were tested using the Kaibab Plateau in Coconino County, Arizona, as a study site. The plateau is isolated from surrounding areas by the Grand Canyon to the south and the Vermillion Cliffs to the north serving as major biogeographical barriers. It is hypothesized that tiger salamanders are not dispersing into or out of the Kaibab Plateau due to geological restrictions. Bd, therefore, should not be present on salamanders on the Kaibab Plateau due to geological restriction. Tiger salamanders in stock tanks located on the Kaibab as well as preserved museum specimens housed in the Arizona State University Natural History Collection were sampled. The results indicate that Bd occurs at low levels on Kaibab Plateau tiger salamanders.
ContributorsOtsuru, Shinji Author (Author) / Collins, James P. (Thesis advisor) / Davidson, Elizabeth (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2019