Innovative restoration strategies are critically needed in the South Texas Plains for controlling increased Opuntia cacti invasions. Using a replicated and randomized experimental study, I have examined the effects of fire seasonality and herbivory on the dominant cacti and herbaceous plant species in this semi-arid ecosystem. Results from this study demonstrate that the combination of fire and wildlife herbivory significantly reduces Opuntia cactus cover. I was able to empirically demonstrate that prescribed fire decreases prickly pear cactus cover. Moreover, this decrease is further exacerbated by the effects of large mammalian herbivores consuming and/or disturbing recently burned mottes. In the absence of fire, both mottes with and without herbivore exclosures increased in size. The ecological insights gained from this study will contribute to the development of management strategies of Opuntia cacti, while promoting the restoration and long-term sustainability of Texas rangelands.

Nutrient availability and herbivory can regulate primary production in ecosystems, but little is known about how, or whether, they may interact with one another. Here I investigate how nitrogen availability and insect herbivory interact to alter above- and belowground plant community biomass in an old-field ecosystem. In 2004, 36 experimental plots were established in which soil nitrogen (N) availability (at three levels) was manipulated and insect abundance (at two levels) in a completely randomized plot design. In 2009, after six years of treatment, I measured aboveground biomass and assessed root production at peak growth. Overall, I found a significant effect of soil N availability on both above- and belowground plant biomass while insects affected only aboveground biomass of subdominant plant species and coarse root production; there were no statistical interactions between N availability and insect herbivory for any response variable. Specifically, responses of aboveground and belowground community biomass to nutrients were driven by reductions in soil N, but not additions, indicating that soil N may not be primarily limiting production in this ecosystem. Insect herbivory altered the aboveground biomass of the subdominant plant species and altered allocation patterns to coarse root production belowground. Overall, the results of six years of nutrient amendments and insect removals suggest strong bottom-up influences on total plant community productivity.

Based on dental, cranial, and postcranial anatomy, members of the Permo-Carboniferous clade Diadectidae are generally regarded as the earliest tetrapods capable of processing high-fiber plant material; presented here is a review of diadectid morphology, phylogeny, taxonomy, and paleozoogeography. Phylogenetic analyses support the monophyly of Diadectidae within Diadectomorpha, the sister-group to Amniota, with Limnoscelis as the sister-taxon to Tseajaia + Diadectidae. Analysis of diadectid interrelationships of all known taxa for which adequate specimens and information are known—the first of its kind conducted—positions Ambedus pusillus as the sister-taxon to all other forms, with Diadectes sanmiguelensis, Orobates pabsti, Desmatodon hesperis, Diadectes absitus, and (Diadectes sideropelicus + Diadectes tenuitectes + Diasparactus zenos) representing progressively more derived taxa in a series of nested clades. In light of these results, it is recommended herein that the species Diadectes sanmiguelensis be referred to the new genus Oradectes, Diadectes absitus be referred to the new genus Silvadectes, and Diasparactus be synonymized with Diadectes to produce Diadectes zenos. The phylogenetic hypothesis also reveals an evolutionary history leading to more efficient oral processing within the lineage, with successive nodes characterized by features indicative of a high-fiber diet. Within Diadectomorpha, diadectids constitute the majority of the species, suggesting that the advent of herbivory resulted in a relatively rapid radiation of species within the group, producing a clade that is markedly more species-rich than other, non-herbivorous diadectomorph taxa. An extensive review of Permo-Carboniferous tetrapod-bearing localities does, however, indicate that diadectids were not a key component of the fauna, discovered at fewer than 50 percent of the sites reviewed. These results counter suggestions that the evolution of Diadectidae led to the formation of the modern terrestrial ecosystem—where a large crop of herbivores supports a much smaller number of carnivores—during the Late Carboniferous and Early Permian.

As sessile organisms, plants are unable to escape from attack by herbivorous insects. To cope with this pressure, plants have evolved several defense strategies, including the production of secondary metabolites, specialized chemicals with ecological functions. Most studies have focused on the role of secondary metabolites in plant defense against chewing insects. Little is known about what compounds are present in phloem sap and how they affect phloem feeding insects. Therefore, I investigated the effects of phenolic compounds on phloem feeders, using Chaitophorus aphids in bioassays with wildtype and transgenic poplar overexpressing the transcription factor MYB 134, which results in elevated levels of tannins and reduced levels of phenolic glycosides. Aphids produced significantly more offspring on MYB 134 plants but showed a significant preference for lower tannin leaf tissue. Analysis of poplar phloem exudates and aphid extracts provides direct evidence that the phenolic glycosides salicin, salicortin and tremulacin are present in poplar phloem and are ingested by aphids. These results are discussed in relation to what is driving the differences in aphid fecundity and choice between plant types. / Graduate / 0309 / 0307 / 0329 / alphareghanwong@gmail.com

Based on dental, cranial, and postcranial anatomy, members of the Permo-Carboniferous clade Diadectidae are generally regarded as the earliest tetrapods capable of processing high-fiber plant material; presented here is a review of diadectid morphology, phylogeny, taxonomy, and paleozoogeography. Phylogenetic analyses support the monophyly of Diadectidae within Diadectomorpha, the sister-group to Amniota, with Limnoscelis as the sister-taxon to Tseajaia + Diadectidae. Analysis of diadectid interrelationships of all known taxa for which adequate specimens and information are known—the first of its kind conducted—positions Ambedus pusillus as the sister-taxon to all other forms, with Diadectes sanmiguelensis, Orobates pabsti, Desmatodon hesperis, Diadectes absitus, and (Diadectes sideropelicus + Diadectes tenuitectes + Diasparactus zenos) representing progressively more derived taxa in a series of nested clades. In light of these results, it is recommended herein that the species Diadectes sanmiguelensis be referred to the new genus Oradectes, Diadectes absitus be referred to the new genus Silvadectes, and Diasparactus be synonymized with Diadectes to produce Diadectes zenos. The phylogenetic hypothesis also reveals an evolutionary history leading to more efficient oral processing within the lineage, with successive nodes characterized by features indicative of a high-fiber diet. Within Diadectomorpha, diadectids constitute the majority of the species, suggesting that the advent of herbivory resulted in a relatively rapid radiation of species within the group, producing a clade that is markedly more species-rich than other, non-herbivorous diadectomorph taxa. An extensive review of Permo-Carboniferous tetrapod-bearing localities does, however, indicate that diadectids were not a key component of the fauna, discovered at fewer than 50 percent of the sites reviewed. These results counter suggestions that the evolution of Diadectidae led to the formation of the modern terrestrial ecosystem—where a large crop of herbivores supports a much smaller number of carnivores—during the Late Carboniferous and Early Permian.

The studies described in this thesis investigate the role of indirect effects in invasion biology. The Introduction provides a brief overview of indirect effects and an outline of the thesis structure. The role of indirect effects in the context of invasion biology is addressed in an in-depth published literature review that comprises the second chapter, providing a theoretical background for the subsequent empirical studies. Chapters Three to Six are comprised of manuscripts that have been published or are under review or in press, which describe studies that investigate the importance of indirect effects in invasion biology using a model system consisting of the alien Asteraceae Senecio madagascariensis, a closelyrelated native, Senecio pinnatifolius, and the insect species with which they interact. Senecio madagascariensis and S. pinnatifolius occur in a similar geographic range in eastern Australia and these studies were conducted in mixed and pure populations of the two species. The herbivore and floral visitor assemblages of the two Senecio species at seven field sites in South-east Queensland were compared using sweep-net sampling, manual searching and floral visitor observation techniques. The floral visitor assemblages were similar between the two species, comprised largely of species of Syrphidae and the European honeybee, Apis mellifera. Herbivore assemblages, however, were highly variable both between species and between sites, with greater herbivore abundance and diversity recorded on the native S. pinnatifolius than its alien congener. The most commonly recorded herbivores were sap-sucking species such as Myridae. The magpie moth, Nyctemera amica was the most common folivore on both Senecio species and laboratory studies demonstrated a clear preference by ovipositing females and feeding larvae of this species for the native Senecio species, over the alien. Field surveys supported these findings, recording greater leaf damage on the native species than the invader. Herbivory levels were lower, rather than higher, in mixed populations than in pure populations, thus there was no evidence that the presence of one species enhanced herbivory in the other. Field pollination trials were conducted to determine whether competition for pollinators or facilitation of pollination occurred in mixed Senecio populations. The presence of the native S. pinnatifolius affected pollinator visitation rates to the alien Senecio; bee visits to S. madagascariensis were significantly reduced by the presence of S. pinnatifolius, whilst syrphid visits increased. However, altered visitation rates were not reflected in seed set. The presence of the alien species had no impact on pollinator visits to the native. Surprisingly, S. pinnatifolius seed set was higher in mixed populations than in pure populations. This might be due to abiotic factors, lower rates of herbivory at these sites or transfer of pollen between species resulting in the production of hybrid seed (if S. madagascariensis has greater male fitness). Hybridisation in the field was investigated using AFLP techniques. No mature hybrid plants were recorded in mixed populations, but hybrid seeds were produced by both species. Senecio pinnatifolius maternal parents produced higher numbers of hybrid seed than expected based on the relative frequencies of the two species, whilst hybridisation in S. madagascariensis was lower than expected. This may indicate greater male fitness of the invader. A range of complex indirect interactions can occur between invasive and native species, with these interactions having the potential to influence the success or failure of the invader and its impacts on co-occurring natives. The Discussion addresses the findings of the studies described here in the context of invasion biology theory.

Timber harvesting results in patches of regenerating forest that are substantially different from surrounding unharvested stands, and provides an opportunity to investigate the effect of habitat change on forest fauna. In this thesis I used timber harvesting as an experimental treatment to investigate the effect of a changed resource base on the home range and resource selection of the swamp wallaby, Wallabia bicolor. I recorded habitat attributes at unharvested control, recently harvested (<12 months old), 5 year old and 10 year old sites. Initially, harvesting removed almost all above-ground plant biomass, although the nitrogen and water content of grass on recently harvested sites was relatively high. Five years after harvesting, sites were dominated by densely regenerating 1-3 m tall Eucalyptus seedlings. Relative to unharvested sites, there was substantial lateral cover and values of a forage quality index were high. In contrast, 10 year old sites supported dense, closed stands of 3-6 m tall eucalypt regeneration, had a moderate amount of lateral cover and had low values of the forage quality index. (For complete abstract open document)

There are numerous examples of how exotic insect pests and pathogens have altered the dominance of native tree species and the ecological processes and function related to those species. While targeted species may persist in a functionally altered state via vegetative sprouting, the widespread decimation of a species can have dramatic direct and indirect consequences for organisms in multiple trophic levels. Devastation due to alien insect herbivores poses the greatest threat to native insect larvae that specialize on the impacted host species. The loss of pollinators whose larvae feed on impacted species and provide services for native plants may also be a serious but yet undocumented indirect threat of these exotic invasions. The disruption of mutualistic relationships between native species will have negative consequences for those species and could potentially benefit exotic species. In the southeastern US, laurel wilt disease (LWD) is impacting numerous species in the Lauraceae family, with the majority of cases observed on Persea borbonia, a common sub-canopy tree found in many Coastal Plain habitats. This species is also known to be the primary larval host of the palamedes swallowtail (Papilio palamedes). While infection rates and crown dieback are catastrophically high (>90%), basal resprouting is a common response in P. borbonia. The exotic Cinnamomum camphora is the only Lauraceae species that has shown resistance to LWD and could benefit from opportunities to replace P. borbonia and other Lauraceae species threatened by LWD. The primary objectives of this study were four fold: 1) to quantify P. borbonia sprouting responses in the field and greenhouse and determine the effect of P. borbonia removal on the composition and abundance of woody and herbaceous plant species in the understory layer, 2) to test the relative suitability of C. camphora as an alternative larval host for P. palamedes, 3) to determine the reliance of the Platanthera ciliaris on P. palamedes for successful pollination and the relative availability of alternative long-tongued pollinators, and 4) to forecast how disease-induced shifts in the relative abundance of native (P. borbonia) and exotic (C. camphora) fruit may alter patterns of consumption and subsequent dispersal of C. camphora by birds. The field component (Grand Bay National Estuarine Research Reserve (GBNERR), Jackson County, MS) of chapter two involved the removal of P. borbonia main stems to mimic the impacts of LWD which resulted in a significant increase (~50%) in light transmission. All treated individuals produced sprouts and the size and number of sprouts was positively related to initial tree size. Following the removal of P. borbonia from treatment plots, Ilex vomitoria showed the greatest increase in basal area after two years. Both woody seedlings and herbaceous plants showed no significant trends in composition and/or abundance over time. In the greenhouse (Southern Illinois University, Carbondale, IL), the stem and leaf biomass of vegetative sprouts was significantly greater in a high-nutrient treatment. Light treatments had no effect on sprout production. Results from chapter two suggest that the loss of P. borbonia from the canopy layer may have little direct effect on plant community dynamics. In addition, I found that sprout production is vigorous in P. borbonia and the capacity to persist and tolerate future disturbances may be enhanced on more nutrient-rich sites. In chapter three, I used laboratory experiments and field observations to compare larval performance and adult female preference of P. palamedes between C. camphora and P. borbonia foliage. My results indicate moderate survivorship on C. camphora (46%) compared to P. borbonia (87%) and there were no differences in first and fourth instar growth rates between treatments. Fourth instars consumed less C. camphora foliage than P. borbonia, but metabolic efficiency did not differ between treatments. In the field and laboratory, I found no oviposition preference for C. camphora relative to P. borbonia. While females laid eggs on C. camphora during laboratory trials, the same number of eggs was also laid on inanimate objects. I conclude that C. camphora is suitable for larval development but host-switching to this species by P. palamedes will be primarily constrained by oviposition behaviors. In chapter four, I monitored pollinator visitation and measured nectar spur lengths of P. ciliaris flowers and proboscis lengths of its floral visitors (at GBNERR). Papilio palamedes was the primary visitor (44 visits) but Phoebis sennae was also observed (4 visits). There were no significant differences among P. ciliaris nectar spurlength and the proboscis lengths of P. palamedes and P. sennae. Fruit set was 55 ± 10.8% with access to pollinators and 0% on bagged inflorescences (pollinators excluded). Although I found a positive relationship between visitation and inflorescence size, there was no such pattern in fruit set, indicating that fruit set was not limited by pollinator visitation within the range of visitation rates I observed. Phoebis sennae may provide supplemental pollination service but is likely constrained by habitat preferences that do not always overlap with those of P. cilaris. Although additional observations are needed, my results suggest that expected LWD-induced declines of P. palamedes will threaten the reproductive success and persistence of P. ciliaris populations. In chapter five, I investigated redundancy between C. camphora and P. borbonia with respect to fruit characteristics (physical and chemical) and selectivity by frugivorous birds (at GBNERR). Across two winter survey periods I observed fruit removal from artificial infructescences. I manipulated background species upon which displays were hung (Myrica cerifera and Triadica sebifera) and the accessibility of the displays. Using motion-activated cameras I confirmed foraging bouts on both P. borbonia and C. camphora fruits by three bird species (Dumetella carolinensis, Turdus migratorius, and Catharus guttatus). There was no significant difference in selectivity between fruit types during year one of my surveys but there was a significant preference for C. camphora in year two, which coincided with significantly lower mean daily temperatures. Background tree species and accessibility had no apparent effect on fruit preference. Total polyphenols and pulp:seed ratio were significantly higher in C. camphora fruit. I conclude that the fruits of C. camphora and P. borbonia represent nearly substitutable resources for native birds. However, native species may prefer C. camphora fruit in times of energetic stress. The decline of P. borbonia will likely increase the consumption and dispersal of C. camphora fruits. Additional studies are required to determine if such changes could ultimately increase the distribution and abundance of this exotic species. Combined, the chapters of this dissertation present substantial empirical evidence for the potential multi-trophic level impacts of an exotic plant disease. While it remains unclear how dramatic these impacts will be, the approach used here is vital for understanding and mitigating the long-term ecological effects of species/disease invasions.

Tallgrass prairie in North America has been highly reduced and degraded by human activity (e.g. agriculture) and now human facilitated restoration is necessary to preserve and reestablish the biodiversity, structure and function of this system. In historical tallgrass prairie large ungulates (e.g. Bison bison) were keystone species that regulated many ecosystem properties and functions. Today, restored prairie often lacks these historical ungulates and white-tailed deer (Odocoileus virginianus) have largely assumed the role of dominant ungulates in small, tallgrass prairie restorations. Little is known about how white-tailed deer affect the development of plant communities and ecosystem function during the onset of prairie restoration. In June 2012 an agricultural field was restored to native prairie species in Konza Prairie Biological Station (KPBS) near Manhattan, KS. Immediately following seeding, experimental plots were established and fences were constructed in half of the plots to excluded white-tailed deer. From 2012 to 2013 deer browse of forbs, aboveground biomass (total, sown forbs, sown grasses, volunteer forbs and volunteer grasses), light availability at the soil surface, soil nutrients, and plant community composition were measure inside and outside of exclosures. The first year of this study occurred during a severe drought which diminished in year two, presenting the opportunity to examine the interaction of climate and deer browse on restoration. In plots where deer had access, the percentage of forbs browsed ranged from 1.3 to 10.5%. The effect of deer browsing on aboveground biomass varied across years for each category of biomass. Total biomass appeared to be regulated more strongly by deer than climate, as unbrowsed plots produced similar biomass in each year despite major climatic variation, while browsed plots did not follow this trend. Across all sampling periods, deer browsing increased light availability by 20%. In year two inorganic N was 19% lower in browsed plots, though potential net N mineralization did not vary between treatments. Plant communities were significantly different between years and, between browsed and unbrowsed plots as time and browsing affected community composition, diversity and richness. Deer browsing increased diversity and richness by 24% and 22% respectively. Community composition was most greatly affected by browsing in year one corresponding to the highest rates of browsing and greatest differences in aboveground biomass. These results indicate that deer can have substantial effects on the initial establishment of prairie communities as well as resource availability from the onset of restoration.

Doctor of Philosophy / Department of Geography / Douglas Goodin / Quantitative remote sensing provides an effective way of estimating and mapping vegetation characteristics over an extensive area. The spatially explicit distribution of canopy vegetative properties from remote sensing imagery can be further used for studies of spatial patterns and processes in grassland systems. My research focused on remote sensing of grassland vegetation characteristics and its applications to spatial analysis of grassland dynamics involving interactions between pyric herbivory and vegetation heterogeneity. In remote sensing of vegetation characteristics, (1) I estimated the foliar pigments and nutritional elements at the leaf level using hyperspectral data. The foliar pigments, chlorophylls and carotenoids, were retrieved by inverting the physical radiative transfer model, PROSPECT. The nutritional elements were modeled empirically using partial least squares (PLS) regression. Correlations were found between the leaf pigments and nutritional elements. This provided insight into the use of pigment-related vegetation indices as a proxy of the plant nutritional quality. (2) At the canopy level, I assessed the use of the broadband vegetation indices, normalized difference vegetation index (NDVI) and green-red vegetation index (GRVI), in detecting vegetation quantity (LAI) and quality (leaf and canopy chlorophyll concentrations). The relationships between vegetation indices and vegetation characteristics were examined in the physical model, PROSAIL, and validated by a field dataset collected from a tallgrass prairie. NDVI showed high correlations with LAI and canopy chlorophylls. GRVI performed even better than NDVI in estimating LAI. A new index GNVI (green-red normalized vegetation index) that combined NDVI and GRVI was proposed to extract leaf chlorophyll concentration. These findings showed the potential of using broadband vegetation indices from multispectral remote sensors to monitor vegetation quantity and quality over a wide spatial extent. In the spatial analysis, I examined interactions between pyric herbivory and grassland heterogeneity at multiple scales from the remote sensing imagery. (3) At a coarse, watershed level, I evaluated effects of fire and large herbivores on the spatial distribution of canopy nitrogen. It was found that the interactive effects of fire and ungulate grazing were present in the watersheds burnt in spring, where a high level of ungulate grazing reduced vegetation density, but promoted canopy heterogeneity. Two grazer species, bison and cattle, were compared. Differences in the vegetation canopy between sites with bison and cattle were observed, which may be related to differences in the grazing intensity, forage behavior and habitat selection between the two grazer species. (4) At a fine, patch level (30 m), bison forage pattern was examined associated with canopy nitrogen heterogeneity. Bison preference for patches with high canopy nitrogen was evident in May. Later in June – September, bison tended to avoid sites with high canopy nitrogen. Vegetation heterogeneity showed significant influences on bison habitat selection in June. Bison preferred sites with low variance in canopy nitrogen, where the patch types were highly aggregated and equitably proportioned.