Pine cone beetles (Conophthorus sp.) are serious pests of many forest ecosystems since they burrow into pine cone tissues for egg deposition, causing the death of the seeds. Management of these beetles in natural and commercial stands of pines has been problematic due to lack of understanding about species limits and distribution. This study was conducted to investigate the phylogeography and phylogenetics of the genus. Several species represented by disjunct populations appear to be monophyletic including Conophthorus edulis, C. mexicanus, C. coniperda, and C. conicollens, whereas C. ponderosae is polyphyletic with many distinct clades isolated by geography. This study explored whether host use or geography has played a greater role in the diversification of this genus, focusing on the polyphyletic C. ponderosae and the monophyletic C. edulis. In the first study, 751bp of the mtDNA CO1 gene were sequenced to reconstruct a phylogeny of the genus, and the distribution and host use were compared to investigate whether these factors were significantly associated. The second study addressed population structure and possible historical influences on the C. edulis and C. ponderosae populations using a nested clade analysis of the mtDNA haplotypes. Despite potential limitations due to sampling, several conclusions could be drawn. Three separate haplotype networks were found for the C. ponderosae haplotypes, indicating that there have been at least three lineages that have associated with P. ponderosa. Geography was significantly associated with the phylogeny at greater distances (>900km), but host use was not significant. At the species level, association with geography is variable. Population structure for C. ponderosae at the species level is minimal, and suggests that there has not been much time for lineage sorting of the haplotypes based on the nested clade analysis as compared to C. edulis.

Phenotypic plasticity, the ability of a genotype to express different phenotypes in different environments, allows organisms to cope with variation in resources and invade novel environments. Biologists have long been fascinated with the costs and tradeoffs that generate and maintain variation in plasticity, such as possible increases in brain size and delays in reproduction associated with the evolution of learning. However, the costs of plasticity vary: many studies have failed to find costs of plasticity, the degree of costs often vary with the system or environments considered, and many costs of plasticity are variable even within the lifetime of an individual. This research adopts a developmental perspective to predict the degree and incidence of costs of plasticity, using host learning in butterflies as a case study. Learning, a mechanism of plasticity that develops through a trial-and-error sampling process, should result in developmental costs and allocation of energy towards development (at the expense of reproduction). Furthermore, costs of learning should be less pronounced in environments for which organisms have innate biases and for learned traits underlain by short-term memory, relative to long-term memory (which requires more developmental re-structuring). This research found support for all three predictions across three levels of costs: behavioral costs, tissue costs, and fecundity trade-offs. Butterflies exhibited genetic variation in their ability to learn to recognize different colored hosts. Genotypes with higher proxies for long-term memory emerged with relatively larger neural investment and smaller reproductive investment. In contrast to these costs of long-term learning, proxies of short-term learning were only correlated with increased exploration of a range of possible resources (types of non-hosts) early in the host-learning process. Family-level costs of plasticity emerged from the ability to learn to locate a red host, for which butterflies do not have an innate bias. Costs of learning were also induced by learning itself: following exposure to novel (red) host environments, individual butterflies, regardless of genetic background, increased exploratory behavior, increased neural investment, and re-allocated energy away from reproduction towards other functions (e.g., flight). Considering developmental mechanisms helps to predict how costs will influence the evolution of learning and plasticity.

In parasites, there are several examples of changes in genome size linked to a parasitic lifestyle—with some species having greatly reduced or expanded genome sizes relative to free-living non-parasitic relatives. What is unknown is whether there is correlated evolution between genome size and host specialization, and whether there is a generalizable framework in predicting genome size evolution in parasites using their genetic architecture and host use ecology. Here, I tested whether genome size of 96 eukaryotic parasites across a wide variety of taxa correlates with host specialization, quantified by the number and phylogenetic relatedness of host species they parasitize. I did not find that genome size and host specialization shared a correlated phylogenetic history; however, I did find that ectoparasites tended to have larger genomes then endoparasites, and that parasitic fungi had more host species then either animal or protozoan parasites. Although no clear trends in the evolution of genomes and host specificity were observed among parasites, my study was significantly limited by gaps in both genome size and host range availability. Future research should seek to address these gaps, as well as improve taxonomic coverage of data, so that trends in the evolution of parasite genome architecture could be adequately tested and delineated.

For insects that develop inside discrete hosts both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. This dissertation examines the effects of host size, host quality, and intraspecific competition on life history and associated traits of populations of the seed-feeding beetle S. limbatus adapted to different host plants, and quantifies population differences in phenotypic plasticity. Populations of the study correspond to divergent clades of the species phylogeography (Colombia and United States). Clades compared differ genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared in Acacia greggii, the larger host. Two host-plant mediated maternal effects were found: offspring matured sooner, regardless of their rearing host, when their mothers were reared on Pseudosamanea guachapele and females laid larger eggs on Ps. guachapele. These results also show that this species in addition to be a smaller is a low quality host. Females also laid more eggs and sooner on A. greggii than in Ps. guachapele and, laid more eggs on P. guachapele when A. greggii seeds were small than when they were large. Eggs were larger when laid on Ps. guachapele and Parkinsonia florida, two hosts that reduce survivorship in all populations. However, Colombia females laid eggs of similar size on Ps. guachapele and Pa. florida, while USA females laid the largest eggs on Pa. florida. Larger beetles were most affected when larval competition was increased and seed size decreased. The responses of different body sized females were asymmetrical showing significant variation in plasticity. Although differences between populations in growth and life history traits appear to be adaptations to the size and quality of their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity play an important role in the evolution of S. limbatus’ diet breadth. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts but also buffering herbivores from selection post-colonization.

Lytopylus is species-rich genus of Agathidinae (Hymenoptera: Braconidae: Agathidinae), but limited information of the genus has been published. Morphological, molecular, and ecological data were compared to resolve the species limits of Lytopylus reared from caterpillars collected the Area de Conservación Guanacaste (ACG) in Costa Rica. Molecular data were initially used to hypothesize species limits, and morphological and host use data were employed to make a final decision when molecular data was indecisive, e.g., when the genetic difference between species was slight. Thirty-two new species are described with image plates of each species. Phylogenetic analyses of the barcode region of the cytochrome c oxidase subunit I (COI) mitochondrial gene was conducted using Neighbor-Joining (NJ) and the Maximum likelihood (ML) analysis. A concatenated COI+28S dataset was analyzed by ML analysis to elucidate evolutionary patterns in ecological characters.

Thesis (PhD)--Stellenbosch University, 2015. / ENGLISH ABSTRACT: Understanding how divergent selection results in the evolution of reproductive isolation (i.e. speciation) is an important goal in evolutionary biology. Populations of herbivorous insects using different host plant species can experience divergent selection from multiple selective pressures which can rapidly lead to speciation. Restio leafhoppers are a group of herbivorous insect species occurring within the Cape Floristic Region (CFR) of South Africa. They are specialised on different plant species in the Restionaceae family. Throughout my thesis I investigated how bottom-up (i.e. plant chemistry/morphology of host plant species) and top-down (i.e. predation and competition) factors drive specialisation and divergence in restio leafhoppers. I also investigated interspecific competition as an important determinant of restio leafhopper community structure. In chapter 2 I quantified host specificity of restio leafhopper species within a local community for 24 months. I found that restio leafhopper species are highly host specific and potentially synchronised with the growth phases of their host plants. In chapter 3 I used a network metric, modularity, to determine whether host plant partitioning in a restio leafhopper community is non-random (i.e. driven by a deterministic process). This metric allows the identification of the components underlying host plant partitioning (modules). I then performed experiments to show that modules, and therefore host plant partitioning, can mostly be explained by preference and performance relationships (i.e. bottom-up factors). In chapter 4 I used null models to test whether niche partitioning in restio leafhopper communities is a general pattern across the landscape. I found non-random niche partitioning, which results from strong host specificity, in all investigated restio leafhopper communities. In addition, I performed binary host choice experiments in the presence and absence of interspecific competition, but found no evidence that interspecific competition narrows host preferences. These findings suggest that host specificity, the cause of niche partitioning, is likely shaped over evolutionary time. Sampling multiple interaction networks across the CFR, in chapter 5, I tested whether restio leafhopper populations are more host specific in species rich communities and regions in the CFR than in species poor communities and regions. I found no positive relationship between restio leafhopper species richness and host specificity at any scale. These findings suggest that specialisation is not driven by interspecific competition. In chapter 6 I investigated host shifts in Cephalelus uncinatus. C. uncinatus has a broader distribution than any single restio species that it can use; suggesting that host plant related divergence may result from geographic range expansion. I found that allopatric and parapatric populations, but not sympatric individuals, using different host plants have divergent host preferences. I also found evidence for morphological divergence in traits related to predator avoidance in population pairs that exhibit divergent host preferences. My findings emphasise the importance of both bottom-up and top-down factors, with the exception of interspecific competition, as determinants of specialisation and divergence in restio leafhoppers. I find no evidence that interspecific competition is an important force structuring restio leafhopper communities. Instead, strongly niche partitioned community structure appears to emerge from the speciation process. / AFRIKAANSE OPSOMMING: Die wyse waarop uiteenlopende seleksie lei tot die evolusie van seksuele isolasie (n.l. spesiasie) is ‘n belangrike vraag in evolutionêre biologie. Plantetende insekpopulasieses wat verskillende gasheerplante gebruik kan onder uinteenlopende veelvoudige seleksie wees en vinnig spesiasie ondergaan. Restio-blaarspringers is ‘n groep plantetende insekspesies wat gespesialiseerd is op verskillende plantspesies in die restio familie. In my tesis ondersoek ek die onder-op (n.l. plantchemie en morfologie) en bo-af seleksiekragte (n.l. predasie en kompetisie) wat lei tot gasheerspesialisasie en -spesiasie in restio-blaarspringers. Ek ondersoek ook die belangrikheid van tussen-spesieskompetisie in gemeenskapsorganisasie. In hoofstuk 2 het ek gasheerspesialisasie gekwantifiseer in ‘n klein restio-blaarspringergemeenskap oor 24 maande. Ek het gevind dat restio-blaarspringers hoogs gasheerspesifiek is en moontlik met die groeifase van hul hoofgasheerplante gesinchroniseerd is. Ek het in hoofstuk 3 ‘n netwerkmetriek, modulariteit, gebruik om te bepaal of restio-blaarspringers se gasheerverdeling nie-stokasties is (n.l. deur deterministiese prosesse veroorsaak is). Hierdie metriek laat ‘n mens toe om die komponente van gasheerverdeling (modules) te identifiseer. Deur middel van eksperimente het ek bepaal dat modules, en dus gasheerverdeling, deur gasheervoorkeur en prestasie (onder-op prossesse) verduidelik kan word. In hoofstuk 4 het ek ondersoek of gasheerverdeling algemeen is. Ek het deur middel van nulmodelle gewys dat gasheerverdeling algemeen is en veroorsaak is deur sterk gasheerspesifiekheid. Ek het ook voorkeureksperimente uitgevoer in die teenwoordigheid en afwesigheid van tussen-spesies kompetisie. Hier het ek geen teken gevind dat huidige tussen-spesies kompetisie gasheervoorkeur beïnvloed nie. My bevindinge in hierdie hoofstuk stel dus voor dat spesialisasie, die oorsaak van gasheerverdeling, oor evolutionêre tyd gevorm word. In hoofstuk 5 het ek ondersoek of populasies van restio-blaarspringers meer gasheerspesifiek is in restio-blaarspringerspesies ryke gemeenskappe en streke as populasies in spesies-arm gemeenskappe en streke. Ek het geen positiewe korrelasie tussen spesiesrykheid en gasheerspesialisasie gevind nie. Dit dui daarop aan dat gasheerspesialisasie, en dus gasheerverdeling, nie deur tussen-spesies kompetisie veroorsaak word nie. In hoofstuk 6 het ek gasheerplantgekoppelde divergensie ondersoek in Cephalelus uncinatus. C. uncinatus se verspreiding is breër as enige restio-spesies wat dit kan gebruik. Dit stel voor dat verspreidingvergroting gasheerverskuiwing mag veroorsaak. Ek het gevind dat populasies wat verskillende plante in allopatrie en parapatrie gebruik uiteenlopende gasheerkeuses maak, maar insekte wat verskillende plante in sympatrie gebruik wys nie ontwrigtende gasheerkeuses nie. Die populasies wat uiteenlopende gasheerkeuses getoon het, het ook verskillende morfologiese teen-predasie eienskappe getoon. Dit dui daarop aan dat predasie belangrik mag wees vir spesiasie in restio-blaarspringers. Die bevindinge van my tesis dui daarop aan dat beide onder-op en bo-af seleksie belangrik is vir gasheer-spesialisasie in divergensie. Nietemin, tussen-spesies kompetisie is nie ‘n belangrike bron vir spesialisasie, divergensie of gemeenskapsorganisasie nie. Gemeenskapsorganisasie is klaarblyklik slegs ‘n gevolg van die spesiasie proses.