Development and Evolution of the Membracid Pronotum

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A major goal of biological studies is to understand how complex forms develop and evolve. Each form is the result of molecular developmental patterning, growth, and the accumulation of changes in these processes from internal and external perturbations in ancestral forms. This dissertation uses each of these lenses to investigate the complex forms in the insect family Membracidae, which arises from the pronotum. In most insects, the pronotum is a simple, domed structure just behind the head, but in membracids it has enlarged and elaborated to look like thorns, plant stipules, fungi, and ants, among other shapes. To investigate this diversity, I rely on landmark based geometric morphometrics to quantify pronotal shapes. The specimens I used included those from a laboratory colony, those collected in and around San Jose, Costa Rica, and those from the Smithsonian National Museum of Natural History collection. In Chapter 1, I examined 5th instar morphogenesis to elucidate the timing of developmental events during the transition from juvenile to adult. These findings informed Chapter 2, which revealed developmental mechanisms related to growth led to transcriptional similarity between the pronotum and wings. In Chapter 3, I identified ontogenetic changes in the patterning of membracid pronotal shape compared to that of a closely related outgroup. Finally, in Chapter 4, I used a phylogenetic framework to investigate developmental modules and the co-occurrence of pronotal shape with two life history characteristics.






Kudla, Anna Marie (2023). Development and Evolution of the Membracid Pronotum. Dissertation, Duke University. Retrieved from


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