From masks to muscles: Mapping facial structure of Nycticebus.

Facial musculature in mammals underlies mastication and nonverbal communicative facial displays. Our understanding of primate facial expression comes primarily from haplorrhines (monkeys and apes), while our understanding of strepsirrhine (lemurs and lorises) facial expression remains incomplete. We examined the facial muscles of six specimens from three Nycticebus species (Nycticebus coucang, Nycticebus javanicus, and Nycticebus menagensis) using traditional dissection methodology and novel three-dimensional facial scanning to produce a detailed facial muscle map, and compared these results to another nocturnal strepsirrhine genus, the greater bushbaby (Otolemur spp.). We observed 19 muscles with no differences among Nycticebus specimens. A total of 17 muscles were observed in both Nycticebus and Otolemur, with little difference in attachment and function but some difference in directionality of movement. In the oral region, we note the presence of the depressor anguli oris, which has been reported in other primate species but is absent in Otolemur. The remaining muscle is a previously undescribed constrictor nasalis muscle located on the lateral nasal alar region, likely responsible for constriction of the nares. We propose this newly described muscle may relate to vomeronasal organ functioning and the importance of the use of nasal musculature in olfactory communication. We discuss how this combined methodology enabled imaging of small complex muscles. We further discuss how the facial anatomy of Nycticebus spp. relates to their unique physiology and behavioral ecology.