Geography and Genetics of Ecological Speciation in Pea Aphids

During ecological speciation, divergent natural selection drives evolution of ecological specialization and genetic differentiation of populations on alternate environments. Populations diverging onto the same alternate environments may be geographically widespread, so that divergence may occur at an array of locations simultaneously. Spatial variation in the process of divergence may produce a pattern of differences in divergence among locations called the Geographic Mosaic of Divergence. Diverging populations may vary in their degree of genetic differentiation and ecological specialization among locations. My dissertation examines the pattern and evolutionary processes of divergence in pea aphids (Acyrthosiphon pisum) on alfalfa (Medicago sativa) and clover (Trifolium pretense). In Chapter One, I examined differences among North American aphid populations in genetic differentiation at nuclear, sequence-based markers and in ecological specialization, measured as aphid fecundity on each host plant. In the East, aphids showed high host-plant associated ecological specialization and high genetic differentiation. In the West, aphids from clover were genetically indistinguishable from aphids on alfalfa, and aphids from clover were less specialized. Thus, the pattern of divergence differed among locations, suggesting a Geographic Mosaic of Divergence. In Chapter Two, I examined genomic heterogeneity in divergence in aphids on alfalfa and clover across North America using amplified fragment length polymorphisms (AFLPs). The degree of genetic differentiation varied greatly among markers, suggesting that divergent natural selection drives aphid divergence in all geographic locations. Three of the same genetic markers were identified as evolving under divergent selection in the eastern and western regions, and additional divergent markers were identified in the East. In Chapter Three, I investigated population structure of aphids in North America, France, and Sweden using AFLPs. Aphids on the same host plant were genetically similar across many parts of their range, so the evolution of host plant specialization does not appear to have occurred independently in every location. While aphids on alfalfa and clover were genetically differentiated in most locations, aphids from alfalfa and clover were genetically similar in both western North America and Sweden. High gene flow from alfalfa onto clover may constrain divergence in these locations.

SOCIAL STRUCTURE OF ASIAN ELEPHANTS (ELEPHAS MAXIMUS) IN SRI LANKA

Asian elephants (Elephas maximus) are critically endangered and live in fragmented populations spread across 13 countries. Yet in comparison to the African savannah elephant (Loxodonta africana), relatively little is known about the social structure of wild Asian elephants because the species is mostly found in low visibility habitat. A better understanding of Asian elephant social structure is critical to mitigate human-elephant conflicts that arise due to increasing human encroachments into elephant habitats. In this dissertation, I examined the social structure of Asian elephants at three sites: Yala, Udawalawe, and Minneriya National Parks in Sri Lanka, where the presence of large open areas and high elephant densities are conducive to behavioral observations. First, I found that the size of groups observed at georeferenced locations was affected by forage availability and distance to water, and the effects of these environmental factors on group size depended on site. Second, I discovered that while populations at different sites differed in the prevalence of weak associations among individuals, a core social structure of individuals sharing strong bonds and organized into highly independent clusters was present across sites. Finally, I showed that the core social structure preserved across sites was typically composed of adult females associating with each other and with other age-sex classes. In addition, I showed that females are social at all life stages, whereas males gradually transition from living in a group to a more solitary lifestyle. Taking into consideration these elements of Asian elephant social structure will help conservation biologists develop effective management strategies that account for both human needs and the socio-ecology of the elephants.

Parsing, Generation and Grammar

Humans use their grammatical knowledge in more than one way. On one hand, they use it to understand what others say. On the other hand, they use it to say what they want to convey to others (or to themselves). In either case, they need to assemble the structure of sentences in a systematic fashion, in accordance with the grammar of their language. Despite the fact that the structures that comprehenders and speakers assemble are systematic in an identical fashion (i.e., obey the same grammatical constraints), the two ‘modes’ of assembling sentence structures might or might not be performed by the same cognitive mechanisms. Currently, the field of psycholinguistics implicitly adopts the position that they are supported by different cognitive mechanisms, as evident from the fact that most psycholinguistic models seek to explain either comprehension or production phenomena. The potential existence of two independent cognitive systems underlying linguistic performance doubles the problem of linking the theory of linguistic knowledge and the theory of linguistic performance, making the integration of linguistics and psycholinguistic harder. This thesis thus aims to unify the structure building system in comprehension, i.e., parser, and the structure building system in production, i.e., generator, into one, so that the linking theory between knowledge and performance can also be unified into one. I will discuss and unify both existing and new data pertaining to how structures are assembled in understanding and speaking, and attempt to show that the unification between parsing and generation is at least a plausible research enterprise. In Chapter 1, I will discuss the previous and current views on how parsing and generation are related to each other. I will outline the challenges for the current view that the parser and the generator are the same cognitive mechanism. This single system view is discussed and evaluated in the rest of the chapters. In Chapter 2, I will present new experimental evidence suggesting that the grain size of the pre-compiled structural units (henceforth simply structural units) is rather small, contrary to some models of sentence production. In particular, I will show that the internal structure of the verb phrase in a ditransitive sentence (e.g., The chef is donating the book to the monk) is not specified at the onset of speech, but is specified before the first internal argument (the book) needs to be uttered. I will also show that this timing of structural processes with respect to the verb phrase structure is earlier than the lexical processes of verb internal arguments. These two results in concert show that the size of structure building units in sentence production is rather small, contrary to some models of sentence production, yet structural processes still precede lexical processes. I argue that this view of generation resembles the widely accepted model of parsing that utilizes both top-down and bottom-up structure building procedures. In Chapter 3, I will present new experimental evidence suggesting that the structural representation strongly constrains the subsequent lexical processes. In particular, I will show that conceptually similar lexical items interfere with each other only when they share the same syntactic category in sentence production. The mechanism that I call syntactic gating, will be proposed, and this mechanism characterizes how the structural and lexical processes interact in generation. I will present two Event Related Potential (ERP) experiments that show that the lexical retrieval in (predictive) comprehension is also constrained by syntactic categories. I will argue that the syntactic gating mechanism is operative both in parsing and generation, and that the interaction between structural and lexical processes in both parsing and generation can be characterized in the same fashion. In Chapter 4, I will present a series of experiments examining the timing at which verbs’ lexical representations are planned in sentence production. It will be shown that verbs are planned before the articulation of their internal arguments, regardless of the target language (Japanese or English) and regardless of the sentence type (active object-initial sentence in Japanese, passive sentences in English, and unaccusative sentences in English). I will discuss how this result sheds light on the notion of incrementality in generation. In Chapter 5, I will synthesize the experimental findings presented in this thesis and in previous research to address the challenges to the single system view I outlined in Chapter 1. I will then conclude by presenting a preliminary single system model that can potentially capture both the key sentence comprehension and sentence production data without assuming distinct mechanisms for each.