Mechanical properties of food and masticatory behavior in llamas, Llama glama

Emily M. Nett, Brielle Jaglowski, Luca J. Ravosa, Dominick D. Ravosa, Matthew J. Ravosa

Research output: Contribution to journalArticlepeer-review


Mammals typically process food items more extensively in their oral cavities than do other vertebrates. Dental morphology, jaw-muscle activity patterns, mandibular movements, and tongue manipulation work to facilitate oral fragmentation of dietary items. While processing mechanically challenging foods, mammals modulate mandibular movements and bite forces via recruitment of greater jaw-adductor muscle forces and protracted biting or chewing. Because jaw-loading patterns are influenced by magnitude; frequency; and duration of muscular, bite, and reaction forces during routine feeding behaviors, relatively larger jaws are thought to be more characteristic of mammals that experience higher masticatory loads due to the processing of mechanically challenging foods. The ease of food fracture during post-canine biting and chewing is mainly determined by food stiffness and toughness. Such foods have been associated with increased loading magnitude and/or greater amounts of cyclical loading (i.e., chewing duration). Dietary properties are thought to modulate cyclical loading through changes in chewing frequency and chewing investment. On the other hand, chewing frequency has been found to be independent of dietary properties in rabbits and primates; however, little evidence exists regarding the influence of dietary properties on these parameters in a broader range of mammals. Here, we assessed chewing behavior in seven adult llamas (Llama glama) processing foods with a wide range of mechanical properties (grain, hay, carrots, and dried corn). Each subject was filmed at 60 frames/s, with video slowed for frame-by-frame computer analysis to obtain length of feeding bout and number of chewing cycles for each food type. These parameters were used to calculate chewing frequency (chews/s), chewing investment (chews/g), and chewing duration (s/g). Chewing frequency was not significantly related to mechanical properties of food, but chewing investment and chewing duration were significantly related to dietary stiffness and toughness. Therefore, cyclical loading is positively influenced by stiff and tough foods. This suggests that variation in jaw morphology in extinct and extant mammals is positively related to dietary stiffness and toughness, which requires greater chewing investment and increased chewing duration.

Original languageEnglish (US)
Pages (from-to)1375-1389
Number of pages15
JournalJournal of mammalogy
Issue number5
StatePublished - Oct 1 2021
Externally publishedYes


  • camelids
  • chewing duration
  • chewing frequency
  • chewing investment
  • cyclical loading
  • mechanical properties of food

ASJC Scopus subject areas

  • Ecology, Evolution, Behavior and Systematics
  • Ecology
  • Animal Science and Zoology
  • Genetics
  • Nature and Landscape Conservation


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