TL;DR: The youngest, most concordant 238U-206Pb zircon age from ash below the fish bed of 413.7±4.4 Ma is reported, contrary to the previously accepted mid-Silurian age based on palynomorphs from adjacent exposures.
Abstract: The oldest-known air-breathing land animal is the millipede Pneumodesmus newmani, found in the Cowie Harbour Fish Bed at Stonehaven, Scotland. Here we report the youngest, most concordant 238U-206Pb zircon age from ash below the fish bed of 413.7±4.4 Ma (±2σ), whereas the youngest age from a tuffaceous sandstone above the fish bed is statistically indistinguishable at 414.3±7.1 Ma. The Cowie Harbour Fish Bed thus appears to be lowermost Devonian (Lochkovian), contrary to the previously accepted mid-Silurian age based on palynomorphs from adjacent exposures. This has implications for the evolutionary timetable of land colonization, as the Cowie ages overlap late Lochkovian zircon ages reported elsewhere for andesite below the nearby (~50 mi) Rhynie Chert, which has more advanced terrestrial biota. The results postdate the possible late Silurian Ludford Lane locality in Shropshire, England. Pneumodesmus newmani is thus not the earliest air-breathing land animal, unless the Ludford Lane locality is younger than presently assigned.
TL;DR: The oldest known evidence of spiracles is demonstrated in Pneumodesmus newmani, proving that the oldest known millipedes were fully terrestrial.
Abstract: New millipede specimens from the Paleozoic of Scotland are described, including Archidesmus macnicoli Peach, 1882, from the Lower Devonian (Lochkovian) Tillywhandland Quarry SSSI and three new taxa—Albadesmus almondi, Pneumodesmus newmani, and Cowiedesmus eroticopodus—from the mid Silurian (late Wenlock—early Ludlow) Cowie Formation at Cowie Harbour. Cowiedesmus eroticopodus new species is placed within the new Cowiedesmidae within the new order Cowiedesmida. Kampecaris tuberculata Brade-Birks from the Lower Devonian (Siegenian) of the Lanark Basin near Dunure is shown not to be a kampecarid myriapod, redescribed as Palaeodesmus tuberculata and placed order incertae sedis within Archipolypoda. Anthracodesmus macconochiei Peach is also redescribed and tentatively placed order incertae sedis within Archipolypoda. Archidesmus macnicoli, Albadesmus almondi, and Palaeodesmus tuberculata are each demonstrated to have broad sternites with laterally placed coxal sockets and paramedian pores containing pa...