Subchondral Bone Apparent Density and Locomotor Behavior in Extant Primates and Subfossil Lemurs Hadropithecus and Pachylemur

Abstract

We analyze patterns of subchondral bone apparent density in the distal femur of extant primates to reconstruct differences in knee posture, discriminate among extant species with different locomotor preferences, and investigate the knee postures used by subfossil lemur species Hadropithecus stenognathus and Pachylemur insignis. We obtained computed tomographic scans for 164 femora belonging to 39 primate species. We grouped species by locomotor preference into knuckle-walking, arboreal quadruped, terrestrial quadruped, quadrupedal leaper, suspensory and vertical clinging, and leaping categories. We reconstructed knee posture using an experimentally validated procedure of determining the anterior extent of the region of maximal subchondral bone apparent density on a median slice through the medial femoral condyle. We compared subchondral apparent density magnitudes between subfossil and extant specimens to ensure that fossils did not display substantial mineralization or degradation. Subfossil and extant specimens were found to have similar magnitudes of subchondral apparent density, thereby permitting comparisons of the density patterns. We observed significant differences in the position of maximum subchondral apparent density between leaping and nonleaping extant primates, with leaping primates appearing to use much more flexed knee postures than nonleaping species. The anterior placement of the regions of maximum subchondral bone apparent density in the subfossil specimens of Hadropithecus and Pachylemur suggests that both species differed from leaping primates and included in their broad range of knee postures rather extended postures. For Hadropithecus, this result is consistent with other evidence for terrestrial locomotion. Pachylemur, reconstructed on the basis of other evidence as a committed arboreal quadruped, likely employed extended knee postures in other activities such as hindlimb suspension, in addition to occasional terrestrial locomotion.

Appendix

Phylogeny in Newick format

((((Nycticebus:35,Perodicticus:35):5,(Otolemur:20,Galago:20):20):33.7,(((((Eulemur fulvus:9.2,Eulemur rubriventer:9.2):10.1,Hapalemur:19.3):4.1,Varecia:23.4):15.9,((Indri:29,Propithecus:29):6.9,Lepilemur: 35.9):3.4):26.9,Daubentonia:66.2):7.5):1.5,((((Alouatta belzebuth: 4.1,A.fusca:4.1):9.5,Ateles:13.6): 6.5,((Saguinus:8.1,Leontopithecus:8.1):2,Callithrix: 10.1):10):19.9,((((((Macaca mulatta:1.7,M. fascicularis:1.7):1.1,M.nemestrina:2.8):5,(Cercocebus: 6.5,Mandrillus:6.5):1.3):1,((Miopithecus: 4,Chlorocebus:4):1,(Cercopithecus diana:5,Cercopithecus l’hoesti:5):0):3.3):4,(((Colobus guereza:0.1,C. polykomos:0.1):1.9,Piliocolobus badius:2): 9,(((Trachypithecus cristata:1,t. vetulus:1,T. phayrei:1): 1,Presbytis hosei:2):2.5,Nasalis:4.5):6.5):2):2,(((Pan:7.7,Gorilla:7.7):8.3,Pongo:16):3,Hylobates: 19):6):15):35.2);