title: Osteological histology of the Pan-Alcidae (Aves, Charadriiformes): correlates of wing-propelled diving and flightlessness.
abstract: Although studies of osteological morphology, gross myology, myological histology, neuroanatomy, and wing-scaling have all documented anatomical modifications associated with wing-propelled diving, the osteohistological study of this highly derived method of locomotion has been limited to penguins. Herein we present the first osteohistological study of the derived forelimbs and hind limbs of wing-propelled diving Pan-Alcidae (Aves, Charadriiformes). In addition to detailing differences between wing-propelled diving charadriiforms and nondiving charadriiforms, microstructural modifications to the humeri, ulnae and femora of extinct flightless pan-alcids are contrasted with those of volant alcids. Histological thin-sections of four species of pan-alcids (Alca torda, †Alca grandis, †Pinguinus impennis, †Mancalla cedrosensis) and one outgroup charadriiform (Stercorarius longicaudus) were compared. The forelimb bones of wing-propelled diving charadriiforms were found to have significantly thicker (∼22%) cortical bone walls. Additionally, as in penguins, the forelimbs of flightless pan-alcids are found to be osteosclerotic. However, unlike the pattern documented in penguins that display thickened cortices in both forelimbs and hind limbs, the forelimb and hind limb elements of pan-alcids display contrasting microstructural morphologies with thickened forelimb cortices and relatively thinner femoral cortices. Additionally, the identification of medullary bone in the sampled †Pinguinus impennis specimen suggests that further osteohistological investigation could provide an answer to longstanding questions regarding sexual dimorphism of Great Auks. Finally, these results suggest that it is possible to discern volant from flightless wing-propelled divers from fragmentary fossil remains.
authors.affiliation: Department of Geological Sciences, Jackson School of Geosciences, University of Texas at Austin, 1 University Station C1100, Austin, Texas; National Evolutionary Synthesis Center, Durham, North Carolina; Research and Collections, North Carolina Museum of Natural Sciences, Raleigh, North Carolina.
authors.forename: N Adam
authors.initials: NA
authors.lastname: Smith
authors.forename: Julia A
authors.initials: JA
authors.lastname: Clarke
mesh_terms: D000818:Animals; D001842:Bone and Bones / Q000033:anatomy & histology / Q000166:cytology / Q000502:physiology; D046091:Charadriiformes / Q000033:anatomy & histology*; D005260:Female; D005426:Flight, Animal / Q000502:physiology*; D005552:Forelimb / Q000033:anatomy & histology / Q000166:cytology / Q000502:physiology; D006614:Hindlimb / Q000033:anatomy & histology / Q000166:cytology / Q000502:physiology; D008297:Male; D059166:Osteology; D010802:Phylogeny; D012737:Sex Factors; D014921:Wings, Animal / Q000033:anatomy & histology* / Q000166:cytology / Q000502:physiology*
keywords: Great Auk; Haversian remodeling; avian cortical bone; bone microstructure; flightlessness; medullary bone; osteosclerosis; pachyostosis