Thesis (Ph. D.)--University of Washington, 1993. / Vita. Includes bibliographical references (leaves [134]-148).

abstract: 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. / Dissertation/Thesis / Doctoral Dissertation Biology 2014

abstract: The Modern Synthesis embodies a theory of natural selection where selection is to be fundamentally understood in terms of measures of fitness and the covariance of reproductive success and trait or character variables. Whether made explicit or left implicit, the notion that selection requires that some trait variable cause reproductive success has been deemphasized in our modern understanding of exactly what selection amounts to. The dissertation seeks to advance a theory of natural selection that is fundamentally causal. By focusing on the causal nature of natural selection (rather than on fitness or statistical formulae), certain conceptual and methodological problems are seen in a new, clarifying light and avenues toward new, interesting solutions to those problems are illustrated. First, the dissertation offers an update to explicitly causal theories of when exactly a trait counts as an adaptation upon fixation in a population and draws out theoretical and practical implications for evolutionary biology. Second, I examine a case of a novel character that evolves by niche construction and argue that it evolves by selection for it and consider implications for understanding adaptations and drift. The third contribution of the dissertation is an argument for the importance of defining group selection causally and an argument against model pluralism in the levels of selection debate. Fourth, the dissertation makes a methodological contribution. I offer the first steps toward an explicitly causal methodology for inferring the causes of selection—something often required in addition to inferring the causes of reproductive success. The concluding chapter summarizes the work and discusses potential paths for future work. / Dissertation/Thesis / Doctoral Dissertation Biology 2016

Once prevalent on the landscape, early-successional habitats are now increasingly threatened in the northeastern United States. As a result, many species that rely on or require habitats dominated by shrubs, young trees, grasses, and forbs have experienced precipitous population declines, leading many organizations to list shrubland habitats and constituent wildlife as a conservation priority. Even-aged forest management (e.g. clearcutting) has been shown to be an efficient and effective means for creating early-successional habitat for certain taxa such as shrubland birds, but it is unfeasible in many situations in southern New England due to public opinion and increased parcelization. Group selection harvests create shrubland conditions in the form of relatively small forest openings (< 1 ha); however, limited attention has been directed toward understanding the extent to which these methods contribute to conservation of early-successional species. In order to assess the conservation value of small forest openings for wildlife in southern New England, I studied two distinct communities associated with early-successional habitats, shrubland birds and bees (Hymenoptera: Apiformes), in openings created by group selection harvests and patch-cutting. In 2014 and 2015, birds were surveyed in small forest openings in western Massachusetts in an effort to describe relationships between species occurrence and patch area as well as other microhabitat-, patch-, and landscape-level variables. Black-and-white warblers, common yellowthroats, chestnut-sided warblers, eastern towhees, and gray catbirds were likely to be present in openings at least 0.3 ha in size, while indigo buntings and prairie warblers had minimum area requirements of 0.55 and 1.07 ha, respectively. Variables within microhabitat-, patch-, and landscape-levels were important for predicting species occurrence. Most notably, prairie warblers were more likely to occur in openings closer to large patches of habitat such as powerline corridors, even if those openings were small in size. I conclude that, despite their inability to support the entire community of shrubland birds in this region, small forest openings can provide habitat for several species of conservation concern if proper attention is given to promoting suitable microhabitat, patch, and landscape characteristics. Bees were sampled in openings as well as adjacent mature forest in an effort to describe the bee community, identify environmental variables influencing bee abundance and diversity, and examine the extent to which openings created by forest management may support bees, as well as potentially augment bee populations within adjacent unmanaged forest. Bees were significantly more abundant and diverse in forest openings than mature forest, but species composition was indistinguishable between openings and forest. Abundance and diversity displayed no relationship with opening size in either openings or forest, but were generally positively related to the amount of early-successional habitat on the landscape. Vegetation characteristics within openings were important in shaping bee communities in openings, with abundance and diversity decreasing with vegetation height and increasing with floral abundance. Notably, eusocial, soft-wood-nesting, and small bees exhibited the opposite pattern in adjacent forest, increasing with the succession of openings and decreasing with greater floral abundance within openings. These results suggest that the creation of small forest openings may help to promote bees both in openings and adjacent mature forest, but certain guilds may be negatively affected.

Imagine a school where everyone tries to help the weakest student, and there is no competition among them except for being nice and supportive! Or companies who, instead of competing with their customers, try to make the world a better place! Yes, it is too big of a goal, but every big move starts from a small step. The small step in my thesis is working on the question "how to make groups work together in a better way" applied to the world of artificial intelligence. To train virtual agents to successfully work together, one must overcome the same problems humans face, such as selfishness. While agents might not be selfish, we typically optimize or preferentially select them on their individual performance. Thus, the selfish agents would also win again. To study and remedy this problem, I made a game for multiple artificial agents competing with each other. I also devised different objective functions so that selfishness might not be the optimal strategy. Specifically, Darwin's theory of Evolution and the group selection mechanism provides a different mechanism to incentivize groups to cooperate. The idea is that when evolving agents under these conditions, the last generation will include only cooperating agents. I used a framework that includes an evolvable neural network for implementing the agents and a genetic algorithm working based on specific rewarding schemes. I found that different rewarding schemes result in different agent behavior. Specifically, when agents evolve in groups whose performance is measured by their weakest member, they not only evolve to become an effective group, but they receive rewards more equally.

In evolutionary theory the existence of self-sacrificing cooperative traits poses a problem that has engendered decades of debate. The principal theories of the evolution of altruism are inclusive fitness, reciprocal altruism, and multilevel selection. To provide a framework for the unification o f these apparently disparate theories, this dissertation identifies two fundamental conditions required for the evolution of altruism: 1) non-zero-sum fitness benefits for cooperation and 2) positive assortment among altruistic behaviors. I demonstrate the underlying similarities in these three theories in the following two ways. First, I show that the game-theoretic model of the prisoner’s dilemm a (PD) is inherent to all three theories. While the PD has been used extensively to model reciprocal altruism, I demonstrate that the n-player PD captures fundamental aspects o f multilevel selection and inclusive fitness in that NPD model parameters relate simply to Simpson’s paradox, the Price covariance equation, and Hamilton’s rule. The tension between hierarchical levels that defines a PD reflects the tension between Abstract levels o f selection that is explicit in multilevel selection theory, and im plicit in the other two theories. Second, Ham ilton’s rule from inclusive fitness theory applies to the other theories. As mentioned, I demonstrate that this rule relates to multilevel selection via the NPD. I also show that Queller’s generalization of Hamilton’s rule applies to the conditional strategies of reciprocal altmism. This challenges the selfish-gene viewpoint by highlighting the fact that it is the phenotypes o f others, not their genotypes, that is critical to the evolution o f altruism. I integrate the PD and H am ilton’s rule as follows: the evolution o f altruism in general involves PD situations in which Hamilton’s rule specifies the necessary relationship between 1) the degree of non-zero-sumness within the PD and 2) the degree of positive assortment among altruistic behaviors. Additional contributions of this research include a demonstration that randomly formed associations can provide the necessary positive assortment for strong altruism to evolve, the development of a new selection decomposition that is symmetrical to the Price equation, and a game-theoretic analysis showing the essential similarity of weak and strong altruism under selection.

No description available.

The Group Selection Questionnaire (GSQ; Cox et al., 2004) is a measure that has been developed to facilitate clinical decisions about a client's readiness for group psychotherapy. The GSQ has demonstrated an ability to predict which clients will experience a reduction in distress through the use of group psychotherapy. This dissertation examines the Group Selection Questionnaire's ability to measure client characteristics that predict the client's ability to benefit from receiving group psychotherapy compared to the ability to benefit from receiving another form of treatment, such as individual or a combination of individual and group psychotherapy, as measured by improved scores on the Outcome Questionnaire (OQ-45; Lambert, Gregersen, Burlingame, & Maruish, 2004). Archival data was analyzed using scores from a sample of 156 college-age participants. Multiple regressions showed that the GSQ and its subscales were effective at predicting improvement in symptomatic distress, but did not demonstrate an ability to predict who would benefit more from group, compared to individual or mixed modalities. Limitations of the study, implications for the measure, and future research are discussed.

Harvesting practices in the southern Appalachians have moved away from clearcutting in favor of variable retention harvesting systems. A study was initiated in 1995-8 to investigate the effects of retaining varying numbers of residual trees on regeneration in seven silvicultural treatments. A second study specifically focused on stump sprouting in only three of those treatments. The treatments for first study included: a clearcut, commercial harvest, leave-tree, shelterwood, group selection, midstory treatment, and an uncut control. The second only focused on the clearcut, leave-tree, and shelterwood. These treatments were implemented in seven stands in Virginia and West Virginia over two physiographic provinces, the Appalachian plateau and Ridge and Valley. The stands were even-aged oak dominated Appalachian hardwood stands on fair quality sites with average ages ranging from 63 to 100 yrs. Permanent plots were randomly located in each stand and all overstory trees (>5m tall) were inventoried and tagged prior to harvest. Regeneration was also quantified. Harvest occurred between 1995-8. For the current studies the plots were re-inventoried 9-11 years post-harvest and all regeneration in all treatments as well as stump sprouts in the selected treatments were quantified. The first study utilized a mixed model ANOVA to analyze five species groups: oak, maple, black cherry-yellow-poplar, miscellaneous, and midstory. Response variables included importance value, average height, and density compared within species group and among treatments. Differences between sprout and seedling origin regeneration were also investigated within species group among treatment. Results indicated that oak densities were similar in all of the treatments, and stump sprouts were larger and more frequent than seedlings. Maple exhibited an increase from pre-harvest overstory importance and exhibited competitive sprouting. The black cherry-yellow-poplar group had few but highly competitive sprouts and a considerable increase in seedling origin regeneration in all treatments. The miscellaneous species densities increased as well with more competitive sprouting in some treatments. The midstory species were excluded from the analysis as it was assumed these species would not occupy canopy positions in a mature stand. The second study investigated differences in the percent of stumps that sprouted and the number of sprouts per stump. The percent data were analyzed using a non-parametric one-way ANOVA and regression analysis, while the sprouts per stump data were compared in a mixed model ANOVA and regression. Species were combined into six groups: the red oak group, chestnut oak, red maple, white oak/hickory group, mixed mesic group, and midstory group. The plateau tended to have reduced sprouting compared to the Ridge and Valley for most species groups and treatments. The red oak group, chestnut oak, and red maple exhibited reduced sprouting with increased residual basal area. The mixed mesic group did not show any effect in sprouting related to residual basal area. Only chestnut oak showed fewer sprouts per stump as residual basal area increased. / Master of Science

The effects of seven silvicultural prescriptions were examined one full growing season post-treatment on five sites in the Ridge and Valley, Cumberland Plateau, and Allegheny Plateau of Virginia and West Virginia. Prescriptions were (1) control, (2) understory control with herbicide, (3) group selection, (4) high-leave shelterwood, (5) low-leave shelterwood, (6) leave tree, and (7) clearcut. The effects of each treatment on the vascular plant community were characterized by changes in species diversity, species composition, exotic species richness, Raunkiaer's life-forms, and growth-forms. Pre-treatment, baseline data, and a randomized block design ensured the precision of post-treatment comparisons. High-disturbance treatments (i.e., low-leave shelterwood, leave tree, clearcut) exhibited lower tree-stratum species richness than unharvested treatments. Prescriptions had no effect on shrub-stratum richness. Compared to the control, woody, herb-stratum richness was higher in harvested treatments. Non-woody herb-stratum richness was unaffected. No differences in herb-stratum species diversity were detected using Shannon's (exp H') or Simpson's (1/D) indices. Non-woody species richness per 2-ha treatment plot was significantly higher than the control for high-disturbance treatments. Pre- vs. post-treatment species composition was least similar for high-disturbance treatments, as measured by qualitative and quantitative community similarity indices. Exotic woody species richness was unaffected by treatment; however, non-woody exotic species were more prevalent in high-disturbance treatments. Hemicrytophytes replaced phanerophytes as the most important life-form for harvested treatments. Therophytes were more important in harvested treatments, while the proportion of cryptophtes and chamaephytes did not change in response to treatments. Tree and shrub growth-forms were less dominant in harvested treatments, while graminoids and annual/biennial forbs were more important for high-disturbance treatments. These data represent only the initial results of a long-term study designed to examine plant-community response to silviculture for one harvest rotation length. Long-term results will aid in devising management strategies that address concerns for biodiversity without devaluing traditional resource demands. / Master of Science