Author ORCID Identifier

Date Available


Year of Publication


Degree Name

Doctor of Philosophy (PhD)

Document Type

Doctoral Dissertation


Arts and Sciences



First Advisor

Dr. Mary A. Arthur


Elevated atmospheric CO2 has been implicated as a driver of increased liana abundance worldwide. Known as disturbance creators and beneficiaries, lianas possess the potential to significantly influence forest ecosystems. I investigated the early-invasion dynamics of Euonymus fortunei (wintercreeper), an evergreen liana that is invading forests in eastern North America, disrupting native plant communities and ecosystem functions.

Wintercreeper is widely cultivated as an ornamental groundcover, frequently invading natural areas via asexual stem growth. Invasion of remote natural areas is dependent upon seed transport and may occur less frequently. I examined the mechanisms of seed dormancy by conducting a ‘move-along’ experiment using fresh and after-ripened seeds. Additionally, I sought to characterize the nature of seed dispersal by birds by deploying seed traps within an invaded forest in central Kentucky. Wintercreeper seeds displayed conditional nondeep physiological dormancy. Although germination occurs at high velocity following cold stratification, a cold period was only facultative to break dormancy. While fresh seeds had greater germination rates (98.6%) compared to after-ripened seeds (85.7%), after-ripened seeds experienced earlier germination (41 days across all treatments). My findings also indicate that bird-mediated seed dispersal occurs throughout the winter from seeds that after-ripen on maternal liana phanerophytes. Overall, these data suggest wintercreeper seeds are capable of dispersing and recruiting in areas with mild to non-existent winters.

The invasion of natural areas by exotic invaders is regulated by biotic and abiotic processes, which influence the invader’s success or failure. I studied the vegetation and soil effects of paired invaded (INV), uninvaded (NAT), and ‘restored’ (RES) sites on the germination and survival of wintercreeper seeds and seedlings. The effect of aril (with vs. without) was also tested, both in the field and in vitro. In the field, total germination and first year survival were 55.6% and 24.2%, respectively, across treatments. Total germination was unaffected by treatments, yet vegetation (P = 0.0016) and aril (P = 0.001) treatments significantly influenced germination rates over time, including delayed germination of seeds with arils. The proportion of germinated seedlings that survived was significantly different based on vegetation (P = 0.054) and aril (P = 0.071) treatments after the first winter of growth, but not prior to the first winter. The proportion of seedling survival was significantly lower among seeds dispersed with an aril and seeds sown within INV treatments; there were no interactive effects.

Finally, I examined how seedling density and growth habit (horizontal vs. vertical) influenced plant survival, growth, and allometry. I also tested extractable soil C, N, P, K, Ca, and Mg prior to planting and after 17 months of soil conditioning. I found evidence that increased planting density negatively influenced growth among individual plants (lower survival, basal diameter, shoot mass, root mass). At the plot level, high-density plantings yielded greater stem length, and shoot, root, and combined biomass, indicating positive frequency dependence for this species. Soil analyses indicated C, N, P, Ca, and Mg significantly increased (P < 0.05) over the course of the experiment. I conclude that in wintercreeper, prominent asexual propagation leads to aggregate populations whose total contributions to above- and belowground biomass are positively correlated with density, even though individual plant fitness is not.

Altogether, this work provides insight into how wintercreeper invasions occur at broad and fine scales. This information will provide a foundation for future wintercreeper studies and aid land managers in their prevention and control strategies.

Digital Object Identifier (DOI)