Ophiacodon long bone histology: the earliest occurrence of FLB in the mammalian stem lineage

The origin of mammalian endothermy has long been held to reside within the early therapsid groups. However, shared histological characteristics have been observed in the bone matrix and vascularity between Ophiacodontidae and the later therapsids (Synapsida). Historically, this coincidence has been explained as simply a reflection of the presumed aquatic lifestyle of Ophiacodon or even a sign of immaturity. Here we show, by histologically sampling an ontogenetic series of Ophiacodon humeri, as well as additional material, the existence of true fibrolamellar bone in the postcranial bones of a member of ‘Pelycosauria’. Our findings have reaffirmed what previous studies first described as fast growing tissue, and by proxy, have disproven that the highly vascularized cortex is simply a reflection of young age. This tissue demonstrates the classic histological characteristics of true fibrolamellar bone (FLB). The cortex consists of primary osteons in a woven bone matrix and remains highly vascularized throughout ontogeny providing evidence to fast skeletal growth. Overall, the FLB tissue we have described in Ophiacodon is more derived or “mammal-like” in terms of the osteonal development, bone matrix, and skeletal growth then what has been described thus far for any other pelycosaur taxa. Ophioacodon bone histology does not show well-developed Haversian tissue. With regards to the histological record, our results remain inconclusive as to the preferred ecology of Ophiacodon, but support the growing evidence for an aquatic lifestyle. Our findings have set the evolutionary origins of modern mammalian endothermy and high skeletal growth rates back approximately 20 M.Y. to the Early Permian, and by phylogenetic extension perhaps the Late Carboniferous.