Biological and Physical Factors Affecting the Natural History and Evolution of Encapsulated Development

The evolution of reproductive strategies involves a complex calculus of costs and benefits to both parents and offspring. Many marine animals produce embryos packaged in tough egg capsules or gelatinous egg masses attached to benthic surfaces. While these egg structures can protect against environmental stresses, the packaging is energetically costly for parents to produce. In this series of studies, I examined a variety of ecological factors affecting the evolution of benthic development as a life history strategy. I used marine gastropods as my model system because they are incredibly diverse and abundant worldwide, and they exhibit a variety of reproductive and developmental strategies.

The first study examines predation on benthic egg masses. I investigated: 1) behavioral mechanisms of predation when embryos are targeted (rather than the whole egg mass); 2) the specific role of gelatinous matrix in predation. I hypothesized that gelatinous matrix does not facilitate predation. One study system was the sea slug Olea hansineensis, an obligate egg mass predator, feeding on the sea slug Haminoea vesicula. Olea fed intensely and efficiently on individual Haminoea embryos inside egg masses but showed no response to live embryos removed from gel, suggesting that gelatinous matrix enables predation. This may be due to mechanical support of the feeding predator by the matrix. However, Haminoea egg masses outnumber Olea by two orders of magnitude in the field, and each egg mass can contain many tens of thousands of embryos, so predation pressure on individuals is likely not strong. The second system involved the snail Nassarius vibex, a non-obligate egg mass predator, feeding on the polychaete worm Clymenella mucosa. Gel neither inhibits nor promotes embryo predation for Nassarius, but because it cannot target individual embryos inside an egg mass, its feeding is slow and inefficient, and feeding rates in the field are quite low. However, snails that compete with Nassarius for scavenged food have not been seen to eat egg masses in the field, leaving Nassarius free to exploit the resource. Overall, egg mass predation in these two systems likely benefits the predators much more than it negatively affects the prey. Thus, selection for environmentally protective aspects of egg mass production may be much stronger than selection for defense against predation.

In the second study, I examined desiccation resistance in intertidal egg masses made by Haminoea vesicula, which preferentially attaches its flat, ribbon-shaped egg masses to submerged substrata. Egg masses occasionally detach and become stranded on exposed sand at low tide. Unlike adults, the encased embryos cannot avoid desiccation by selectively moving about the habitat, and the egg mass shape has high surface-area-to-volume ratio that should make it prone to drying out. Thus, I hypothesized that the embryos would not survive stranding. I tested this by deploying individual egg masses of two age classes on exposed sand bars for the duration of low tide. After rehydration, embryos midway through development showed higher rates of survival than newly-laid embryos, though for both stages survival rates over 25% were frequently observed. Laboratory desiccation trials showed that >75% survival is possible in an egg mass that has lost 65% of its water weight, and some survival (<25%) was observed even after 83% water weight lost. Although many surviving embryos in both experiments showed damage, these data demonstrate that egg mass stranding is not necessarily fatal to embryos. They may be able to survive a far greater range of conditions than they normally encounter, compensating for their lack of ability to move. Also, desiccation tolerance of embryos may reduce pressure on parents to find optimal laying substrata.

The third study takes a big-picture approach to investigating the evolution of different developmental strategies in cone snails, the largest genus of marine invertebrates. Cone snail species hatch out of their capsules as either swimming larvae or non-dispersing forms, and their developmental mode has direct consequences for biogeographic patterns. Variability in life history strategies among taxa may be influenced by biological, environmental, or phylogenetic factors, or a combination of these. While most prior research has examined these factors singularly, my aim was to investigate the effects of a host of intrinsic, extrinsic, and historical factors on two fundamental aspects of life history: egg size and egg number. I used phylogenetic generalized least-squares regression models to examine relationships between these two egg traits and a variety of hypothesized intrinsic and extrinsic variables. Adult shell morphology and spatial variability in productivity and salinity across a species geographic range had the strongest effects on egg diameter and number of eggs per capsule. Phylogeny had no significant influence. Developmental mode in Conus appears to be influenced mostly by species-level adaptations and niche specificity rather than phylogenetic conservatism. Patterns of egg size and egg number appear to reflect energetic tradeoffs with body size and specific morphologies as well as adaptations to variable environments. Overall, this series of studies highlights the importance of organism-scale biotic and abiotic interactions in evolutionary patterns.

Essays on Microeconometrics

My dissertation has three chapters which develop and apply microeconometric tech- niques to empirically relevant problems. All the chapters examines the robustness issues (e.g., measurement error and model misspecification) in the econometric anal- ysis. The first chapter studies the identifying power of an instrumental variable in the nonparametric heterogeneous treatment effect framework when a binary treat- ment variable is mismeasured and endogenous. I characterize the sharp identified set for the local average treatment effect under the following two assumptions: (1) the exclusion restriction of an instrument and (2) deterministic monotonicity of the true treatment variable in the instrument. The identification strategy allows for general measurement error. Notably, (i) the measurement error is nonclassical, (ii) it can be endogenous, and (iii) no assumptions are imposed on the marginal distribution of the measurement error, so that I do not need to assume the accuracy of the measure- ment. Based on the partial identification result, I provide a consistent confidence interval for the local average treatment effect with uniformly valid size control. I also show that the identification strategy can incorporate repeated measurements to narrow the identified set, even if the repeated measurements themselves are endoge- nous. Using the the National Longitudinal Study of the High School Class of 1972, I demonstrate that my new methodology can produce nontrivial bounds for the return to college attendance when attendance is mismeasured and endogenous.

The second chapter, which is a part of a coauthored project with Federico Bugni, considers the problem of inference in dynamic discrete choice problems when the structural model is locally misspecified. We consider two popular classes of estimators for dynamic discrete choice models: K-step maximum likelihood estimators (K-ML) and K-step minimum distance estimators (K-MD), where K denotes the number of policy iterations employed in the estimation problem. These estimator classes include popular estimators such as Rust (1987)’s nested fixed point estimator, Hotz and Miller (1993)’s conditional choice probability estimator, Aguirregabiria and Mira (2002)’s nested algorithm estimator, and Pesendorfer and Schmidt-Dengler (2008)’s least squares estimator. We derive and compare the asymptotic distributions of K- ML and K-MD estimators when the model is arbitrarily locally misspecified and we obtain three main results. In the absence of misspecification, Aguirregabiria and Mira (2002) show that all K-ML estimators are asymptotically equivalent regardless of the choice of K. Our first result shows that this finding extends to a locally misspecified model, regardless of the degree of local misspecification. As a second result, we show that an analogous result holds for all K-MD estimators, i.e., all K- MD estimator are asymptotically equivalent regardless of the choice of K. Our third and final result is to compare K-MD and K-ML estimators in terms of asymptotic mean squared error. Under local misspecification, the optimally weighted K-MD estimator depends on the unknown asymptotic bias and is no longer feasible. In turn, feasible K-MD estimators could have an asymptotic mean squared error that is higher or lower than that of the K-ML estimators. To demonstrate the relevance of our asymptotic analysis, we illustrate our findings using in a simulation exercise based on a misspecified version of Rust (1987) bus engine problem.

The last chapter investigates the causal effect of the Omnibus Budget Reconcil- iation Act of 1993, which caused the biggest change to the EITC in its history, on unemployment and labor force participation among single mothers. Unemployment and labor force participation are difficult to define for a few reasons, for example, be- cause of marginally attached workers. Instead of searching for the unique definition for each of these two concepts, this chapter bounds unemployment and labor force participation by observable variables and, as a result, considers various competing definitions of these two concepts simultaneously. This bounding strategy leads to partial identification of the treatment effect. The inference results depend on the construction of the bounds, but they imply positive effect on labor force participa- tion and negligible effect on unemployment. The results imply that the difference- in-difference result based on the BLS definition of unemployment can be misleading

due to misclassification of unemployment.

Multilevel Governance and Accountability: Does Decentralization Promote Good Governance?

Today, the trend towards decentralization is far-reaching. Proponents of decentralization have argued that decentralization promotes responsive and accountable local government by shortening the distance between local representatives and their constituency. However, in this paper, I focus on the countervailing effect of decentralization on the accountability mechanism, arguing that decentralization, which increases the number of actors eligible for policy making and implementation in governance as a whole, may blur lines of responsibility, thus weakening citizens’ ability to sanction government in election. By using the ordinary least squares (OLS) interaction model based on historical panel data for 78 countries in the 2002 – 2010 period, I test the hypothesis that as the number of government tiers increases, there will be a negative interaction between the number of government tiers and decentralization policies. The regression results show empirical evidence that decentralization policies, having a positive impact on governance under a relatively simple form of multilevel governance, have no more statistically significant effects as the complexity of government structure exceeds a certain degree. In particular, this paper found that the presence of intergovernmental meeting with legally binding authority have a negative impact on governance when the complexity of government structure reaches to the highest level.

Food Swamps, Obesity & Health Zoning Restrictions on Fast Food Restaurants

Protecting public health is the most legitimate use of zoning, and yet there is minimal progress in applying it to the obesity problem. Zoning could potentially be used to address both unhealthy and healthy food retailers, but lack of evidence regarding the impact of zoning and public opinion on zoning changes are barriers to implementing zoning restrictions on fast food on a larger scale. My dissertation addresses these gaps in our understanding of health zoning as a policy option for altering built, food environments.

Chapter 1 examines the relationship between food swamps and obesity and whether spatial mapping might be useful in identifying priority geographic areas for zoning interventions. I employ an instrumental variables (IV) strategy to correct for the endogeneity problems associated with food environments, namely that individuals may self-select into certain neighborhoods and may consider food availability in their decision process. I utilize highway exits as a source of exogenous variation .Using secondary data from the USDA Food Environment Atlas, ordinary least squares (OLS) and IV regression models were employed to analyze cross-sectional associations between local food environments and the prevalence of obesity. I find even after controlling for food desert effects, food swamps have a positive, statistically significant effect on adult obesity rates.

Chapter 2 applies theories of message framing and prospect theory to the emerging discussion around health zoning policies targeting food environments and to explore public opinion toward a list of potential zoning restrictions on fast-food restaurants (beyond moratoriums on new establishments). In order to explore causality, I employ an online survey experiment manipulating exposure to vignettes with different message frames about health zoning restrictions with two national samples of adult Americans age 18 and over (N1=2,768 and N2=3,236). The second sample oversamples Black Americans (N=1,000) and individuals with high school as their highest level of education. Respondents were randomly assigned to one of six conditions where they were primed with different message frames about the benefits of zoning restrictions on fast food retailers. Participants were then asked to indicate their support for six zoning policies on a Likert scale. Subjects also answered questions about their food store access, eating behaviors, health status and perceptions of food stores by type.

I find that a message frame about Nutrition and increasing Equity in the food system was particularly effective at increasing support for health zoning policies targeting fast food outlets across policy categories (Conditional, Youth-related, Performance and Incentive) and across racial groups. This finding is consistent with an influential environmental justice scholar’s description of “injustice frames” as effective in mobilizing supporters around environmental issues (Taylor 2000). I extend this rationale to food environment obesity prevention efforts and identify Nutrition combined with Equity frames as an arguably universal campaign strategy for bolstering public support of zoning restrictions on fast food retailers.

Bridging my findings from both Chapters 1 and 2, using food swamps as a spatial metaphor may work to identify priority areas for policy intervention, but only if there is an equitable distribution of resources and mobilization efforts to improve consumer food environments. If the structural forces which ration access to land-use planning persist (arguably including the media as gatekeepers to information and producers of message frames) disparities in obesity are likely to widen.