In the lower 300 m of the fluvial Willwood Formation of northwest Wyoming, most large concentrations of vertebrate fossils consist of disarticulated and broken skeletal remains that occur in widespread, tubular, thin (2 cm-1 m) greenish- and bluish-gray mudstones that are low in percentage of free iron, aluminum, and manganese, relatively high in percentages of organic carbon, and often show evidence of clay eluviation. These units lie atop sesquioxide-rich, sometimes clay-illuviated variegated unit (red, yellow, purple, and mottled variants) and are relicts of the A horizons of podzolic spodosols (aquods).

The wide distribution of bone fragments, the numerical proportions of bone elements, weathering of the fragments, and biogenic evidence indicate the vertebrate remains accumulated gradually as litter on the surface of the soils and became incorporated into the A horizon. The bones were disarticulated, broken, and otherwise disturbed by the action of scavengers prior to burial. Occurrence of abundant vertebrate remains in discrete, readily identifiable, and widespread paleosol units makes these units valuable biostratigraphic markers (zonules, faunules). Evidence of geologically short-term lag accumulation of the bones is important to assemblage analysis; the paleosol concentrations result from attritional mortalities that more closely reflect composition of the life assemblage than do remains concentrated by fluvial transport or predators.