Insights for modern invasion ecology from biotic changes of the Clarksville Phase of the Richmondian Invasion (Ordovician, Katian)

Abstract. The frequency of biotic invasions in modern ecosystems is increasing due to global trade moving taxa outside their native ranges and climate change facilitating establishment of taxa in previously inhospitable regions. Thus, developing a holistic understanding of biotic invasions and how they impact ecosystems over different timescales—from annual to geologic timescales—is vital. Herein we examine a geologically brief invasion event, the Clarksville Phase of the Richmondian Invasion. Prior analyses have established general ecological and evolutionary patterns across the entire Richmondian Invasion, but recent sequence stratigraphic refinement makes analysis of individual invasion pulses possible for the first time. We examine biotic change across the Clarksville Phase and identify invasion impacts on diversity, paleocommunity composition, and niche stability. Invader arrival and success was strongly linked to increased propagule pressure facilitated by sea-level changes. Invaders initially colonized deep subtidal environments and then moved offshore facilitated by rapid niche evolution during the invasion interval. Invasive taxa that attained the largest population sizes belonged to previously underutilized ecological guilds. Overall, the introduction of the invasive taxa resulted in increased diversity that was maintained into the postinvasion interval accompanied by a change in community composition in which the invaders became dominant paleocommunity members. Combined, these analyses document a biotic invasion facilitated by climate change that increased local diversity through invaders occupying underutilized ecospace and competition-related niche contraction on millennial timescales. Developing a long-term perspective to accompany shorter-term studies facilitates predicting the long-term impacts of modern invasions and creating better-informed policies and practices.