The skull of the Miocene Spheniscus urbinai (Aves, Sphenisciformes): osteology, brain morphology, and the cranial pneumatic systems.

Spheniscus urbinai represents one of four extinct Spheniscus species from the Cenozoic of southern South America, known from several poorly described diversely complete skulls and postcranial elements. Here, we present a review of the cranial osteology of all known specimens (collected in Argentina, Chile, and Peru), including a paleoneurological analysis using CT scans, and an exploration of its cranial pneumaticity compared to other extinct and living seabirds. Our results show that among Spheniscus species, S. urbinai exhibits slightly greater cranial pneumaticity than the living species. Additionally, we confirm previous findings which indicate that the marked reduction of cranial pneumaticity-which is characteristic of living penguins-occurred early during the Eocene (as observed in the Antarctic penguin MLP 12-I-20-1, but not in the coeval Anthropornis).

The Origin and Evolutionary Consequences of Skeletal Traits Shaped by Embryonic Muscular Activity, from Basal Theropods to Modern Birds.

Embryonic muscular activity (EMA) is involved in the development of several distinctive traits of birds. Modern avian diversity and the fossil record of the dinosaur-bird transition allow special insight into their evolution. Traits shaped by EMA result from mechanical forces acting at post-morphogenetic stages, such that genes often play a very indirect role. Their origin seldom suggests direct selection for the trait, but a side-effect of other changes such as musculo-skeletal rearrangements, heterochrony in skeletal maturation, or increased incubation temperature (which increases EMA). EMA-shaped traits like sesamoids may be inconstant, highly conserved, or even disappear and then reappear in evolution. Some sesamoids may become increasingly influenced in evolution by genetic-molecular mechanisms (genetic assimilation). There is also ample evidence of evolutionary transitions from sesamoids to bony eminences at tendon insertion sites, and vice-versa. This can be explained by newfound similarities in the earliest development of both kinds of structures, which suggest these transitions are likely triggered by EMA. Other traits that require EMA for their formation will not necessarily undergo genetic assimilation, but still be conserved over tens and hundreds of millions of years, allowing evolutionary reduction and loss of other skeletal elements. Upon their origin, EMA-shaped traits may not be directly genetic, nor immediately adaptive. Nevertheless, EMA can play a key role in evolutionary innovation, and have consequences for the subsequent direction of evolutionary change. Its role may be more important and ubiquitous than currently suspected.