Diversity partitioning of a Late Ordovician marine biotic invasion: controls on diversity in regional ecosystems

Biotic invasions are a common feature of the fossil record, yet remarkably little is known about them, given their enormous potential to reveal the processes that regulate local and regional diversity over long time scales. We used additive diversity partiticming to examine how diversity structure changed as a result of a marine biotic invasion in tropical, shallow and deep subtidal environments spanning approximately 4 Myr in the Late Ordovician. The biotic invasion increased richness in the regional ecosystem by nearly 40%. Within-habitat turnover diversity accounts for most of the increase in richness, with between-habitat turnover diversity contributing a lesser amount. Increases in these components of diversity were accommodated by increased packing of species along a depth gradient and increased habitat heterogeneity. Diversity metrics that incorporate taxon abundance (Shannon information, Simpson's D) show similar patterns and reveal that many invading taxa were locally abundant and widespread in their occurrence. Extinction of incumbent taxa did not foster the invasion; rather the invasion appears to be linked to a regional or global warming event. Taken together, these observations indicate that these Late Ordovician marine communities were open to invasion and not saturated with species. Moreover, the increase in species diversity caused by the invasion was not ephemeral; instead it lasted for at least 1 Myr. Similar studies of other biotic invasions in the fossil record are necessary to determine (1) the factors, such as extinction of incumbents or resource limitation, that may facilitate or inhibit invasion in ancient ecosystems; (2) how local and regional ecosystems respond to invasion; and (3) the extent to which biotic invasions play a substantial role in ecosystem change through geologic time.