The earliest burst of necrophagous dung beetles in South America revealed by the Cenozoic record of Coprinisphaera

New ichnological, sedimentological and palaeobotanical information from a Miocene palaeosol succession from Patagonia bearing abundant fossil brood balls of dung beetles (Coprinisphaera) allow inferring novel aspects of the evolutionary history, biology and feeding habits of Scarabaeinae, along with the palaeoenvironmental and palaeoclimatic conditions in which they nested. Coprinisphaera tonnii and Coprinisphaera akatanka, both attributed to necrophagous dung beetles, represent 40.2% of the specimens. Considering their scarce or null record in older Cenozoic units from South America, these high values reveal that the first burst of necrophagous Scarabaeinae took place during the Miocene. Some Coprinisphaera preserve characters that indicate the developmental stage of the ball occupant and the adult emergence success. Both ichnospecies show the egg chamber isolated from the provision chamber, which was associated with a higher preference of cleptoparasites for necrophagous balls. The comparison among the abundance of traces of nest intruders in Coprinisphaera attributed both to necrophagous and to coprophagous beetles from different units of South America indicates that the presence of such intruders would be independent of the nature of the organic matter contained within the balls. Phytolith analyses performed in Coprinisphaera and extant necrophagous and coprophagous balls indicate that the comparison between the relative abundance of phytoliths in the wall of the brood ball, their infilling and the bearing palaeosol is a useful tool for inferring the feeding behaviour of the trace makers and support the attribution of Coprinisphaera tonnii and Coprinisphaera akatanka to necrophagous Scarabaeinae. Sedimentary and palaeosol analyses indicate that the beetles would have nested in well-drained soils showing mollic-like features, in grass-dominated habitats, under a seasonal sub-humid, temperate–warm climate.Fil: Cantil, Liliana Fernanda. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Bellosi, Eduardo Sergio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Sánchez, María Victoria. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Gonzalez, Mirta Gladys. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Genise, Jorge Fernando. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentin