Masticatory myology of the llama (Lama glama, Camelidae) and comparisons with other camelids and euungulates

Camelids are the only living representatives of the Suborder Tylopoda, and present a unique set of osteo-myological masticatory features, differing from all other extant euungulates. They combine selenodont dentition and rumination with a fused symphysis, and roughly plesiomorphic muscle proportions. Despite its potential relevance as an euungulate model in comparative anatomy studies, the available data is strikingly scarce. The present study represents the first description of the masticatory muscles of a Lamini, analyzing the functional morphology of Lama glama and other camelids in a comparative framework. Both sides of the head of three adult specimens from Argentinean Puna were dissected. Descriptions, illustrations, muscular maps, and weighing of all masticatory muscles were performed. Some facial muscles are also described. The myology of llamas confirms that camelids possess relatively large temporalis muscles, with Lama being less extreme than Camelus. This plesiomorphic feature is also recorded in suines and some basal euungulates. Conversely, the direction of the fibers of the M. temporalis is mainly horizontal, resembling grinding euungulates such as equids, pecorans, and some derived suines. Although the M. masseter of camelids and equids do not reach the particularly modified configuration of pecorans, in which it is rostrally extended and arranged horizontally, the posterior sectors of Mm. masseter superficialis and pterygoideus medialis have acquired relatively horizontal disposition in the former lineages, suitable for protraction. The pterygoidei complex presents several bundles, and its relative size is intermediate between suines and derived grinding euungulates. The whole masticatory muscles are relatively light when compared to jaw weight. The evolution of the masticatory muscles and chewing of camelids implied that grinding abilities were reached with less extreme modifications of the topography and/or proportions than pecoran ruminants and equids. A relatively large M. temporalis recruited as a powerful retractor during the power stroke is a key feature of camelids. The relaxed pressure on chewing derived from the acquisition of rumination explains the slenderer build masticatory musculature of camelids compared to other euungulates except ruminants.