Ecology & Evolutionary Biology Dissertations and Theses

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  • Publication
    Phylogenomic systematics, bioacoustics, and morphology of frogs from Madagascar reveals that background noise drives the evolution of high frequency acoustic signaling
    (University of Kansas, 2019-12-31) Hutter, Carl; Glor, Richard; Brown, Rafe; Holder, Mark; Cartwright, Paulyn; Macdonald, Stuart
    Madagascar is considered a globally important biodiversity hotspot, having some of the highest rates of species endemism in the world and has been the focus of substantial effort from researchers to understand the evolution of its distinctive biota. This is especially true for frogs, where the microcontinent hosts an impressively diverse amphibian fauna totaling over 500 species, where a large amount of this diversity has only been described relatively recently. Much of this accelerated taxonomic progress can be attributed to the combination of DNA barcoding and the widespread application of bioacoustics enabling more efficient species identification and characterization of new lineages. The availability of bioacoustic data for the majority of species has revealed the incredible diversity of acoustic signals in Malagasy frogs; despite the incredible acoustic diversity in Madagascar, explanations for the evolution of distinct advertisement calls have been little explored. Acoustic signaling is important to frogs because it is the primary mechanism of communication and mate selection and therefore it is expected that acoustic communication and factors that drive variation in acoustic signals should be under strong selection. Frogs from the most species-rich genus Boophis from the family Mantellidae in Madagascar communicate acoustically during a rainy and short breeding season and aggregate around water bodies in high abundance making Boophis an ideal model system to address the evolution of advertisement calls. A potential explanation for signal diversity is the acoustic adaptation hypothesis, which predicts that natural and sexual selection drive optimization of signal transmission and perception across different habitats. In many organisms, the efficiency of acoustic signal transmission can be affected by habitat structure or background noise. One prediction of the acoustic adaptation hypothesis is that environmental ambient noise, which is the background level of sound in the environment, will drive signal evolution if the noise is sufficiently similar and lessens the receiver’s perception of the signal. Because Boophis reproduce near water bodies and most often in stream habitats that might be noisy from the sound of rushing water, they present an excellent system to address acoustic interference. In Chapter 1, I describe a new species of Boophis that is morphologically cryptic with its sister species but differs remarkably in advertisement call, which underlines the importance of call variation in the genus. To contribute to the taxonomic progress and integration of bioacoustic data, I describe a new species of Boophis by using these multiple lines of evidence and also suggesting future research to understand the evolution of advertisement calls in the genus. I also develop the acoustic analysis pipeline used for acquiring frequency traits from thousands of calls rapidly, where these methods will be used in Chapter 5. In this study, I also suggest that reproductive character displacement could be driving divergence in advertisement calls and that other important factors from the environment could lead to broad patterns in the evolution of advertisement calls. In Chapter 2, to understand signal evolution across a broad range of taxa, a strongly supported phylogenetic hypothesis is needed and I estimate a new multi-locus phylogeny using Sanger sequencing. The systematics of frogs from the family Mantellidae have had a long and turbulent history where Mantellidae was considered a family relatively recently. In Boophis tree frogs, early researchers considered them as belonging to the Asian genus Rhacophorus because of their strong similarities and breeding habitat in water bodies. Furthermore, despite the numerous works that contributed to understanding the molecular phylogeny of Boophis many aspects of their evolutionary relationships remain insufficiently supported and a complete Sanger multi-locus phylogeny had not been estimated. For understanding the relationships among Boophis frogs, I estimated a multi-locus phylogeny from eight Sanger markers that includes as much diversity as possible. Despite these efforts, I could not adequately support the phylogenetic relationships among Boophis taxa and adding additional Sanger markers would not be likely to resolve these relationships. In Chapter 3 I aim to expand upon this dataset and develop a new sequencing technology to obtain thousands of markers affordably. The widespread use of high-throughput sequencing technologies to sequence large portions of organisms’ genome has led to new and exciting challenges and questions that can be addressed with the massive increase in sequence data these new methods provide. The objective of sequence capture is to sequence genomic regions typically through hybridization-based capture from a previously designed set of known markers and benefits from the potential to acquire markers that are useful at all evolutionary time-scales. Therefore, I developed a sequence capture probe set called FrogCap to sequence ~15,000 genomic markers that can be used across the entire frog radiation. I compare the efficacy of the probe set on six phylogenetic scales and quantify the number of markers sequenced, depth of coverage, missing data, and parsimony informative sites, and I also compared differences between these measures across different types of data. The results from this chapter show that FrogCap is a very promising new sequence capture probe set that can be used across all frogs. In Chapter 4 I test the effectiveness of FrogCap by addressing the systematics of frogs from the family Mantellidae, by comparing the FrogCap sequence capture results to those from transcriptomes. The phylogeny of Mantellidae remains unresolved and contentious, where the phylogenetic relationships among subfamilies and genera are not well delimited. Prior to this it was thought the different groups in Madagascar were non-monophyletic and were thought to be from several different families. I address the weak phylogenetic support and also test the FrogCap probe set on Mantellidae frogs, comparing the probe set to transcriptomic sequencing of samples from the same groups. I find that both FrogCap and transcriptomes work similarly well for resolving these difficult and contentious relationships. FrogCap sequence capture also provided several advantages over the transcriptomic data; FrogCap sequences non-protein coding markers from across the genome, such that these other types of markers could be useful by providing sequence data from potentially neutrally evolving genomic regions and also can serve as another line of phylogenetic evidence when compared to other data types. Transcriptomes also provide advantages through a larger amount of sequence data and lesser gene discordance because the transcriptomes are much longer and thus providing more resolution than shorter markers. I find that the FrogCap probe set is an effective tool at disentangling difficult phylogenetic problems, and that transcriptomic sequencing is less effective for phylogenetics. In Chapter 5, I estimate a new phylogeny for Boophis tree frogs using the FrogCap probe set and address whether acoustic interference leads to the evolution of higher frequency advertisement calls in Boophis frogs. I integrate an unprecedented dataset incorporating data collected from all previous chapters which includes a massive dataset of 300+ acoustic recordings from nearly every species in the genus, a new Boophis time-calibrated phylogeny using a backbone of ~15,000 genomic markers acquired from the sequence capture data combined with the data from Chapter 2 for full species sampling and used soft tissue computed tomography on 28 species to acquire detailed morphological information from the larynges of males. After finding that the sequence capture dataset provides strong statistical support for nearly every node in the tree, I time-calibrate the phylogeny and test the acoustic adaptation hypothesis. I find that Boophis tree frogs are evolving higher frequency acoustic signals in loud stream habitats, which is supported after correcting for body size. These results are further evidenced by laryngeal measurements from the CT Scans, where I find that loud stream frog laryngeal morphology is decoupled from the predicted relationship to body size that is found in quiet stream frogs.
  • Publication
    Evolution of Sexual Signals in the Drosophila saltans Species Group
    (University of Kansas, 2019-12-31) Colyott, Kaila; Gleason, Jennifer M; Holder, Mark T; Macdonald, Stuart J; Blumenstiel, Justin P; Walters, Jamie R
    The complex courtship signaling of three of the four species groups of the Sophophora subgenus (e.g., melanogaster, obscura, and willistoni) has been studied. In this dissertation, I examined the complex courtship signaling of the fourth species group, saltans, for the first time. In the first chapter, I assessed through what modalities courtship signals were communicated in Drosophila saltans (sensu stricto). No single modality ablation eliminated mating, and all ablations affected mating, thus courtship was multimodal. In the second chapter, I examined if the role of two modalities (i.e., vision and audition) during mating were correlated in nine species of the saltans species group. In particular, I investigated if the two modalities were assessed in an integrated manner (i.e., positively correlated), or if there was a tradeoff between the role of two modalities (i.e., negatively correlated). The role of vision and audition varied from playing no role to being necessary for mating success and were not correlated when considering independent contrasts. In Chapter Two, I also described the auditory courtship signal of each species of the group and found variation in song production with one species producing an elaborated song element and two species producing reduced songs. Furthermore, song production (reduced or elaborated) was not associated with the role of audition in mating success. In the third chapter, I examined if the two song types (i.e., pulse and beep) in the elaborated song of D. sturtevanti communicated one or multiple messages and if the messages were unique or redundant. The song of D. sturtevanti conveyed multiple redundant messages. In summary, I found that courtship signaling was complex by being multimodal in D. saltans (sensu stricto) and containing both multiple, and redundant messages in the auditory modality of D. sturtevanti. Also, I found that apparent signal complexity of song was not associated with the role of song during courtship.
  • Publication
    Redescription of Wushaichthys exquisitus and phylogenetic revision of Thoracopteridae
    (University of Kansas, 2019-12-31) Shen, Chenchen; Beard, K. Christopher; Arratia, Gloria; Schultze, Hans-Peter; Lieberman, Bruce; Smith, W. Leo
    Wushaichthys exquisitus, a small fish from the Middle Triassic of Guizhou Province, China, was named and studied by Xu et al. (2015), and Xu et al. interpreted it as the most primitive species in the family Thoracopteridae because of the presence of the following features: the skull roof is formed by the frontal bone, which is laterally expanded whereas the dermopterotic is expanded posteriorly; the parietal bone is fused with the dermopterotic; the supraorbital sensory canal ends in the frontal; presence of a triangular extrascapular, which is separated from its counterpart medially by the posttemporals; and presence of a narrow and vertical preopercle with an anterior process contacting the maxilla. However, this hypothesis was questioned because W. exquisitus also shares some features with Peltopleuridae, such as the deepened-flank scales, which means W. exquisitus should be a peltopleurid fish. A redescription of Wushaichthys exquisitus is conducted here based on five new specimens. Several new characters are described here, including the vertical preopercle sensory canal, which extends from ventral to dorsal margin of preopercle with a branch to the maxillary process; a small hyomandibular process on the antero-dorsal margin of opercle; and the presence of two lateral lines. A phylogenetic analysis based on 101 characters and 23 taxa was conducted. The results of the phylogenetic analysis suggest that W. exquisitus is the most basal taxon of the extinct family Thoracopteridae, confirming previous studies. The new results also indicate that Thoracopteridae is the sister group of Peltopleuridae, and these two families form the order Peltopleuriformes. This conclusion also agrees with several previous studies. Based on the detailed redescription, the diagnosis of the genus Wushaichthys is emended as well as the diagnoses of Wushaichthy exquisitus and of the family Thoracopteridae.
  • Publication
    Mathematical Modeling of Neisseria meningitidis: A Case Study of Nigeria
    (University of Kansas, 2019-12-31) Ojo, Mayowa Micheal; Orive, Maria E; Agusto, Folashade B; Peterson, A. Townsend; Reuman, Daniel
    Bacterial meningitis remains one of the deadliest infectious diseases in the African meningitis belt. It is defined as an acute inflammation of the meninges, the protective membranes that cover the brain and spinal cord. This deadly disease has different serogroups with geographical distribution and epidemic potential varying amongst each serogroup. To date, six of these serogroups have been identified as causative agents of epidemics. The effect of this disease cannot be ignored due to its high morbidity and mortality; however, vaccination has played a major role in preventing the spread of the disease in the population. The work presented here was therefore designed to study the effect of constant and pulse vaccination in controlling the disease in the population using a non-impulsive model and an impulsive models. The two models were analyzed qualitatively to determine the conditions for eradicating the disease. Numerical simulation of the non-impulsive model showed that meningitis can be effectively controlled in the population using an imperfect vaccine with an efficacy greater than 75% and high vaccine coverage rate of at least 85%. For the impulsive model, we obtain the disease-free periodic solution, and the model is locally asymptotically stable when the threshold quantity, Rp, is less than one. Furthermore, numerical simulations showed that the final infected population size is lower when applying the impulsive vaccination strategy compared with the scenario without vaccination. Thus, the disease burden in the population decreases with increasing vaccination pulses. Lastly, we present a deterministic model to study the dynamics of the co-infection of multiple strains (serogroup A and serogroup C) in the presence of vaccination. For the scenario in which we set the transmission probability of a strain to zero, our results show that the co-infection model exhibits competitive exclusion (a strain driving the other strain into extinction when both are at endemic equilibrium). However, for the scenario in which we set the transmission probability of one strain greater than that of the other (transmission probability of neither strain is equal to zero), we observed a trade-off mechanism that enables co-existence of the two strains. In this case, regardless of the greatest reproduction number value of each strain, the strain with the highest transmission probability in co-infected individuals will dominate in the population without driving the other strain into extinction. We further analyzed and simulated the model without co-infection. The model exhibit competitive exclusion when R0A R0C 1 (strain A drives out strain C) or when R0C R0A 1 (strain C drives out strain A). Additionally, our results show that when the two strains have the same reproduction number at their endemic state, the two strains will co-exist but the dominance of a strain will depend on the initial size of its population.
  • Publication
    On the nature of genetic variation in yellow monkeyflower (Mimulus guttatus): combining theoretical and empirical approaches
    (University of Kansas, 2019-08-31) Brown, Keely Elizabeth; Kelly, John K; Gamblin, Truman C; Orive, Maria E; Sikes, Benjamin A; Hileman, Lena C
    Evolution via natural selection requires standing variation in a population while simultaneously diminishing it at loci under selection, a paradox that continues to vex evolutionary biologists. The work presented here contributes to our understanding of how variation can be maintained by selection, why decreasing genetic variation is detrimental on an individual level, and how genetic variation can have complicated effects on fitness-related phenotypes. Chapter one demonstrates that antagonistic pleiotropy caused by a fitness tradeoff between polygenic traits can maintain variation at a single locus by generating emergent overdominance. The results are surprisingly robust to perturbations in the underlying assumptions about random mating and environmental heterogeneity. Chapter two provides an explanation for how mating system can influence the maintenance of variation. Inbreeding depression in the Iron Mountain population of yellow monkeyflower (Mimulus guttatus) is potentially severe enough to maintain outcrossing and is also predicted by the load of rare alleles carried by a genotype. Chapter three illustrates the complex relationships between genotype and phenotype that are likely underappreciated in typical QTL mapping or GWAS approaches, due to tightly linked antagonistic effectors. It provides a mechanism for studying the influence of genetic variation on complex fitness-related phenotypes, like flower size.
  • Publication
    A revision of the genus Colletes in America north of Mexico (Hymenoptera, Colletidae)
    (University of Kansas, 1952-05-31) Stephen, William Procuronoff
    The bee genus Collates abounds in America north of the tropic of Cancer and is widely distributed throughout the world, with the exception of Australia. This study is limited to those species occurring in America north of Mexico, for material from the Central American countries is meagre and areas of collection disjunct. The work is primarily of a systematic nature and contributes little to an evolutionary or phylogenetic study of the genus. However, in conjunction with Noskiewicz’s (1936) treatise on the Palearctic members of the genus plus a critical treatment of the Central American fauna, it should form a suitable basis tor the inauguration of such a project.
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    A reclassification of Western Hemisphere Typhlocybinae (Homoptera, Cicadellidae)
    (University of Kansas, 1951-05-31) Young, David A., Jr.
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    Speciation and distribution of the crayfishes of the Ozark Plateaus and Ouachita provinces
    (University of Kansas, 1951-05-31) Williams, Austin B.
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    A monograph of the Saldidae (Hemiptera) of North and Central America and the West Indies
    (University of Kansas, 1949-05-31) Hodgden, Burton B.
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    The Pangoniinae (Diptera: Tabanidae) of Nearctic America
    (University of Kansas, 1933-05-31) Brennan, James M.
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  • Publication
    Hydrogeomorphic drivers of functional diversity in riverine fishes at an intercontinental scale
    (University of Kansas, 2019-05-31) Mathews, Gregory; Thorp, James H; Alexander, Helen; Smith, William L
    Functional diversity (FD) has been used as a metric to gauge the health and stability of fish communities in many different environments, but few studies have examined FD on an intercontinental scale. This study formally examined the FD of riverine fishes at a macrosystem scale based on hydrogeomorphic features in similar climates between two continents. To accomplish this goal, I sampled fish in five systems representing three ecoregions (terminal basin, mountain steppe and grasslands) across the United States of America and Mongolia. I then investigated how FD changed between and within each continent and ecoregion type. As expected, FD was strongly correlated with species diversity. Additionally, I found that FD, specifically functional richness, was higher in wider, deeper rivers and decreased with faster, more sloped systems. This suggests that FD increases similarly to species richness as one moves from high elevation headwaters toward larger, lowland systems. However, a community containing redundant species that offer no novel traits to the community can complicate this generality. This project should serve as a complement to earlier work as well as provide a foundation for future studies that attempt to more thoroughly understand functional diversity at a macrosystem scale.
  • Publication
    High supply, high demand: A unique nutrient addition decouples nitrate uptake and metabolism in a large river
    (University of Kansas, 2019-05-31) Kelly, Michelle Catherine; Burgin, Amy J; Husic, Admin; Sikes, Benjamin A
    Our current understanding of the relationship between nitrate (NO3-) uptake and energy cycling in rivers is primarily built on studies conducted in low-nutrient (NO3- < 1 mg-N L-1), small (discharge < 1 m3 s-1) systems. Recent advances in sensor technology have allowed for continuous measures of whole-river NO3- uptake, allowing us to address how the relationship between nutrient uptake and metabolism changes over time and space during a nutrient addition in a large river. We treated a six-month controlled nitrogen (N) waste release into the Kansas River (conducted by the City of Lawrence, KS) as an ecosystem-scale nutrient addition experiment. We deployed four NO3- and dissolved oxygen sensor arrays along a 33 km study reach from February to May 2018 to continuously monitor diel NO3--N and stream metabolism. We then evaluated NO3- uptake using the extrapolated diel method and modeled stream metabolism using the single station method. We found the highest uptake rates closest to the nutrient release point (866 g-N m-2 d1), despite high NO3- supply (4.36 mg-N L-1). Net ecosystem productivity was increasingly autotrophic with distance from the release, with the highest respiration rates observed closest to the release point (7.09 g-O2 m-2 d1). However, uptake was decoupled from metabolism metrics, likely due to fine-scale hydrologic and biotic factors. Overall, our work sheds light on the ability of large rivers to retain and transform nutrients, while demonstrating that the fine-scale mechanisms that regulate nutrient retention in large rivers are still largely unknown.
  • Publication
    Phylogenetic and Biogeographic Analysis of Cretaceous Spatangoid Echinoids from the Western Interior Seaway
    (University of Kansas, 2019-05-31) Byrum, Steven; Lieberman, Bruce; Smith, William L; Beard, Kenneth C
    Members of the echinoid order Spatangoida, a highly diverse and abundant marine invertebrate clade, were important denizens of the Cretaceous Western Interior Seaway (WIS), an epicontinental seaway that divided North America in two during an interval of greenhouse conditions between roughly 100 and 65 million years ago. Despite being well represented in the Cretaceous rocks of Texas and surrounding areas, the phylogenetic patterns of spatangoid echinoids of the WIS, especially at the species level, have yet to be fully resolved. Further, because of their complex morphologies, they represent one of the few marine invertebrate groups from this time period that are amenable to phylogenetic study. Another interesting question pertaining to taxa found in the WIS is their biogeographic origins and whether they represent an endemic radiation or are instead derived from one or more invasions from other regions. In order to reconstruct a hypothesis of spatangoid phylogenetic relationships, a parsimony analysis was conducted using a character matrix of 32 characters collected from 20 species. Species that occur in the WIS were considered comprehensively; species from other regions such as South America, Europe, and North Africa were additionally incorporated into the analysis. Phylogenetic biogeographic analysis was then conducted on the spatangoid phylogeny using a modified version of Fitch Parsimony Analysis, and several episodes of vicariance and range expansion were identified across the phylogeny. These were possibly further related to some of the various major episodes of sea-level rise and fall during the Cretaceous.
  • Publication
    Dancing in the Dark: The evolution of visually mediated courtship behaviors and sexual dimorphisms in spotted winged Drosophila
    (University of Kansas, 2019-05-31) Roy, Paula R; Gleason, Jennifer M; Blumenstiel, Justin; MacDonald, Stuart; Morgan, Theodore; Rivers, Trevor; Walters, James
    I employed a multidisciplinary approach to examine the function and evolution of sexual dimorphisms, male wing spots and male wing displays, across four species subgroups of Drosophila. In chapter one, I explored the function of wing spots in three species of the D. suzukii group using female choice tests by placing mating pairs in darkness. I found that females do not show a statistical preference for spotted males in choice experiments, but that vision is either required for or facilitates mating success. The wing spot may not be as important for mating decisions as previously hypothesized but could enhance courtship displays in more complex environments. In chapter two, I used whole genome sequence data to build a phylogeny of the spotted winged Drosophila in the D. melanogaster group, identified the likelihood of ancestral states of the wing spot character, and tested for conservation of the morphology. The rate of evolution is too rapid to determine if wing spots are ancestral and subsequently gained or lost throughout the subgroups, but multiple convergent events were identified. The wing spot did not have a phylogenetic signal and was correlated with frontal courtship display behaviors, suggesting that wing spots are likely a rapidly fluctuating sexually selected character. In chapter three, I measured visually mediated behaviors of 13 species of spotted and non-spotted Drosophila and mapped behavior to the phylogeny to identify associations between behavior and wing spot morphology that could explain the observed evolutionary patterns. Courtship behavior is labile, and visually mediated long-term copulation acceptance is associated with wing spot morphology. The results in total suggest that species delimitation could be driven by female choice sexual selection on male dimorphisms. In chapter four, I detailed potential functions of the wing spot. This study lays the groundwork for further study on the function of various male displays observed in species where males have sexual dimorphisms.
  • Publication
    Wild bee responses to land use change: investigating the role of bee body size
    (University of Kansas, 2019-05-31) Mayes, Daphne; Smith, Deborah; Alexander, Helen; Sikes, Ben; Hagen, Robert; Brown, J. Christopher
    Land use change impacts biodiversity through many facets including an alteration of habitat and the resources required to sustain species and populations. Wild bees (Hymenoptera: Apoidea) are essential pollinators for many wild and cultivated flowering plants, and exhibit marked differences in life history traits. The consequences of land use change may therefore affect wild bees differently, but this remains poorly understood for many taxa. Body size is a trait that can vary widely across and within bee species, and has important ramifications for several aspects of bee fitness including foraging range. To better understand wild bee responses to land use change, I focused on bee body size from the scale of the community (interspecific) and within a species (intraspecific). In Chapter 1, I examined tallgrass prairie remnants and restorations across eastern Kansas and found that both types of prairie hosted a similar abundance, diversity, and body size of trap-nesting bees over both study years. The bee community composition differed between prairie types in the first study year but not the second. Regardless of prairie type, bee diversity increased with increasing forb diversity but did not have significant associations with landscape composition. Trap-nesting bee abundance and interspecific body size did not vary in relation to local forb diversity or landscape composition. In Chapter 2, I again used the context of the tallgrass prairie to focus on the response (i.e., total offspring produced, intraspecific body size, and sex ratio) of a single bee species, Heriades carinata, to prairie type, forb diversity, and landscape composition. My results indicate that within this species, foraging mothers provision a greater number albeit smaller size of offspring (both male and female) in prairies with increasing forb diversity. These trends were not extended to the landscape scale, however, indicating that the resources immediately surrounding nest sites may have a greater influence on bees in these sites. In Chapter 3, I extend the question of interspecific bee body size to stingless bee communities collected across a deforestation gradient in Rondônia, Brazil. Stingless bees are diverse and important pollinators in tropical systems, but little is known about how they respond to habitat loss and fragmentation from a trait-based perspective. I found that larger bees were collected more often in areas with less forest and within landscapes that had greater isolation between remaining forest patches, while smaller bees were found in areas with a greater amount of forest and shorter distances between forest patches. In Chapter 4, I find that Africanized honey bees were observed more frequently in open (i.e., deforested) areas but still rely on some amount of forest in the landscape. Taken together, my results suggest that body size differently affects wild bee responses to land use change.
  • Publication
    Relationships between bruchid beetles (Amblycerus robiniae) and honey locust trees (Gleditsia triacanthos)
    (University of Kansas, 1971-12-31) Mathwig, John E.
    This study concerns the relationships between the bruchid beetle Amblycerus robiniae and the honey locust tree Gleditsia triacanthos. Since Amblycerus concentrates specifically on Gleditsia, and on specific parts of this tree, it may influence plant density. Thus, such a phytophagous insect can effect and control the species composition and the structure of a plant community.
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    Evolution of novel genetic programs: insights from flower and fruit development
    (University of Kansas, 2019-08-31) Sengupta, Aniket; Hileman, Lena C; Hileman, Lena C.; Cartwright, Paulyn; Mort, Mark; Smith, Wm L; Buechner, Matthew
    An outstanding question in evolutionary biology is how genetic programs (interaction between multiple genes or their products) that define novel phenotypes evolve. There are two major ways such genetic programs that define novel phenotypes can evolve. First, by a de novo assembly of previously non-interacting genes during the origin of the novel phenotype. Secondly, by co-option (re-deployment, re-recruitment) of existing programs from other functions towards defining the novel phenotype. Monosymmetry of flowers is a novel phenotype that has evolved at least 130 times from polysymmetric flowers during the diversification of flowering plants. Flower monosymmetry in the order Lamiales is defined by an interaction of CYCLOIDEA, RADIALIS, DIVARICATA, and DIVARICATA and RADIALIS interacting factors (CYC–RAD–DIV–DRIF). This interaction is best understood in the Lamiales species Antirrhinum majus. The evolutionary history of the CYC–RAD–DIV–DRIF genetic program that defines Lamiales flower monosymmetry is unclear. It is an open question whether this genetic program was assembled de novo near the base of Lamiales during the evolution of flower monosymmetry or it was co-opted from a different function. We find evidence that the CYC–RAD–DIV–DRIF genetic program, which is crucial for defining the novel phenotype of flower monosymmetry in Lamiales, was likely co-opted from a different function, possibly fruit and ovary development. We come to this conclusion through a comparative analysis between representative taxa from the Lamiales and its close relative the Solanales. This suggests that the evolution of flower monosymmetry in Lamiales may have been facilitated by the availability of the CYC–RAD–DIV–DRIF genetic program. This genetic program was likely co-opted in defining flower monosymmetry in Lamiales for the following reasons. This program was ancestrally involved in regulating cell size: and this function makes it a likely candidate for defining a novel phenotype that has variously shaped petals and aborted stamens. Also, the CYC–RAD–DIV–DRIF genetic program functions by a competitive interaction between RAD and DIV proteins—the two competing proteins could easily be co-opted in defining the two morphologically distinct regions of the monosymmetric flowers. Little is known about the regulators that affect the transcription of CYC in Lamiales. We find predictive, bioinformatics-based evidence for a Lamiales-specific transcriptional autoregulation of CYC, suggesting that the evolution of flower monosymmetry in this lineage was associated with an autoregulation-mediated sustained, stable, and high transcription of CYC. Our data elucidates the evolutionary origin of the CYC–RAD–DIV–DRIF genetic program that defines flower monosymmetry in Lamiales. This interaction was likely co-opted en bloc from fruit and ovary development, with additional lineage-specific changes in gene regulation leading to transcriptional autoregulation of CYC. Entangled in this question is the fact that ovaries and fruits of many species in the tribe Antirrhineae (order Lamiales), including Antirrhinum majus, are monosymmetric—with unequal dorsal and ventral locules. We determine the genetic and micromorphological basis of this phenotype and also estimate its evolutionary history. We identify at least five evolutionary transitions from polysymmetric to monosymmetric ovaries, and at least seven reversals to polysymmetry across the tribe. Ovary monosymmetry in Antirrhinum and its closest relatives is likely controlled non-cell autonomously by a CYC–RAD interaction. CYC upregulates RAD expression in dorsal petals and stamens in early stages of development, and RAD protein from these tissues migrates to the dorsal locule to increases cell proliferation in dorsal ovary wall, causing the ovary to be monosymmetric.
  • Publication
    Population ecology of the softshell turtle, Trionyx muticus
    (University of Kansas, 1976-05-31) Plummer, Michael V.
    Nesting success of Trionyx muticus was studied during two field seasons from individually marked nests, Supplementary information was obtained from eggs subjected to submersion experiments in the laboratory, Open areas on high ridges of sandbars are preferred nesting sites; apparently females recognize these areas from the water. Predation on nests was low. The major cause of egg mortality was submersion produced by extended rises in water level; laboratory experiments suggest that egg viability is decreased when submersion lasts for more than one day. Nesting success was lower in 1973, an unusually wet season, than in 1974 which was unusually dry, Evidence indicates that there must be a minimum number of hatchlings in a nest to insure escape from the nest.