1. Temperate deciduous forests in the United States are located in the most densely populated states across the northern and mid-Atlantic east coast. Land development and associated human activities result in small forests that are susceptible to anthropogenic influences, such as urbanization and non-native plant invasion. 2. The overall objective of this study was to assess spatial and temporal drivers of forest vegetation structure and diversity in small forests embedded across developed landscapes. We assessed woody plant composition across spatial gradients (i.e., urbanization and non-native plant invasion) and a temporal gradient (i.e., time since canopy closure) across 38 forests along the east coast of the United States in northern Delaware and southeastern Pennsylvania. 3. Surprisingly, we found the invasion gradient was not related to the urban gradient across our forests. Across all forests, the canopy consisted of native species (98% of all trees), whereas the forest understory was most vulnerable to nonnative plant invasion (65% of all woody plant stems). Greater native species richness in forest canopies and understories with increasing urbanization supports the conclusion that urban forests maintain native species and are not inherently degraded ecosystems. Non-native plant invasion has a strong influence on understory plant communities, and the duration of intact forest canopy had a strong negative correlation with non-native plant invasion suggesting intact forests can resist invasion. 4. Synthesis. This is the first study to compare simultaneously the importance of invasion and urbanization in determining plant community composition in forests embedded across developed landscapes, and to discover that younger forests harbor more invasive plants and urban forests maintain native plants. K E Y W O R D S nonmetric multidimensional scaling, non-native plant invasion, temperate deciduous forest, urban forests, urbanization, woody plant community | 2367
Forest structural diversity and its spatiotemporal variability are constrained by environmental and biological factors, including species pools, climate, land‐use history, and legacies of disturbance regimes. These factors influence forest responses to disturbances and their interactions with structural diversity, potentially creating structurally mediated emergent properties at local to continental spatial scales and over evolutionary time. Here, we present a conceptual framework for exploring the emergent properties that arise from interactions between forest structural diversity and disturbances. We synthesize and present definitions for key terms, including emergent property, disturbance, and resilience, and highlight various types and examples of emergent properties, such as (1) interactions with species composition, (2) interactions with disturbance frequency and intensity, and (3) evolutionary changes to communities. Although emergent properties in forest ecosystems remain poorly understood, we describe a foundation for study and applied management of forest structural diversity to enhance forest restoration and resilience.
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