The rise and fall of late Paleozoic trilobites of the United States

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Abstract
Based on range data and generic composition, four stages of evolution are recognized for late Paleozoic trilobites of the contiguous United States. Stage 1 occurs in the Lower Mississippian (Kinderhookian-Osagean) and is characterized by a generically diverse association of short-ranging, stenotopic species that are strongly provincial. Stage 2 species are present in the Upper Mississippian and consist of a single, eurytopic, pandemic genus, Paladin. Species of Stage 2 are much longer-ranging than those of Stage 1, and some species may have persisted for as long as 12 m.y. Stage 3 is present within Pennsylvanian and Lower Permian strata and consists initially of the eurytopic, endemic genera Sevillia and Ameura as well as the pandemic genus Ditomopyge. During the middle Pennsylvanian the very long-ranging species Ameura missouriensis and Ditomopyge scitula survived for more than 20 m.y. During the late Pennsylvanian and early Permian, a number of pandemic genera appear to have immigrated into what is now North America. Stage 4 is restricted to the Upper Permian (late Leonardian-Guadalupian) strata and is characterized by short-ranging, stenotopic, provincial genera.The main causal factor controlling the four-stage evolution of late Paleozoic trilobites of the United States is interpreted to be eustacy. Whereas Stage 1 represents an adaptive radiation developed during the Lower Mississippian inundation of North America by the Kaskaskia Sequence, Stage 2 is present in strata deposited during the regression of the Kaskaskia sea. Stage 3 was formed during the transgression and stillstand of the Absaroka Sequence and, although initially endemic, Stage 3 faunas are strongly pandemic in the end when oceanic circulation patterns were at a maximum. A mid-Leonardian sea-level drop caused the extinction of Stage 3 fauna. Sea-level rise near the end of the Leonardian and into the Guadalupian created an adaptive radiation of stentopic species of Stage 4 that quickly became extinct with the latest Permian regression.