A b s t r a c t T h e n i n e -b a n d e d a r m a d i l l o , D a s y p u s novemcinctus, is a member of the family Dasypodidae, which contains all extant species of armadillos and represents the most diverse group of xenarthran mammals by their speciation, form, and range of scratch-digging ability. This study aims to identify muscle traits that reflect specialization for fossorial habit by observing forelimb structure in D. novemcinctus and comparing it among armadillos using available myological data. A number of informative traits were observed in D. novemcinctus and among Dasypodidae, including the absence of m. rhomboideus profundus, the variable presence of a m. articularis humeri and m. coracobrachialis, two heads of m. triceps brachii with scapular origin, and a lack of muscle mass devoted to antebrachial supination. Muscle mass and myosin heavy chain (MHC) isoform content were also quantified from our forelimb dissections. New osteological indices are additionally calculated and reported for D. novemcinctus. Collectively, the findings emphasize muscle mass and power output for limb retraction and specialization of the distal limb for sustained purchase of soil by strong pronation and carpal/digital flexion. Moreover, the myological traits assessed here provide a valuable resource for interpretation of muscle architecture specializations among digging mammals and future reassessment of armadillo phylogeny.

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As a muscular hydrostat, the tongue undergoes complex deformations during most oral behaviors, including chewing and drinking. During these behaviors, deformations occur in concert with tongue and jaw movements to position and transport the bolus. Moreover, the various parts of the tongue may move and deform at similar timepoints relative to the gape cycle or they may occur at different timepoints, indicating regional biomechanical and functional variation. The goal of this study is to quantify tongue biomechanics during chewing and drinking in pigs by characterizing intrinsic deformations of the tongue across multiple regions simultaneously. Tongue deformations are generally larger during chewing cycles compared to drinking cycles. Chewing and drinking also differ in the timing of regional length and width, but not total length, deformations. This demonstrates functional differences in the temporal dynamics of localized shape changes whereas the global properties of jaw-tongue coordination are maintained. Finally, differences in the trade-off between length and width deformations demonstrate that the properties of a muscular hydrostat are observed at the whole tongue level, but biomechanical variation (e.g., changes in movements and deformations) at the regional level exists. This study provides new critical insights into the regional contributions to tongue deformations as a basis for future work on multidimensional shape changes in soft tissues.

In mammals, chewing movements can be modified, or flexible, in response to changes in food properties. Variability between and within food in the temporal characteristics of chewing movements can impact chewing frequency and rhythmicity, which in turn may affect food breakdown, energy expenditure and tooth wear. Here, we compared total chewing cycle duration and intra-cycle phase durations in pigs chewing on three foods varying in toughness and stiffness: apples (low toughness, low stiffness), carrots (high toughness, low stiffness), and almonds (high toughness, high stiffness). We also determined whether within-food variability in timing parameters is modified in response to changes in food properties. X-ray Reconstruction Of Moving Morphology (XROMM) demonstrates that the timing of jaw movements are flexible in response to changes in food properties. Within each food, pigs also exhibited flexibility in their ability to vary cycle parameters. The timing of jaw movements during processing of high-toughness foods is more variable, potentially decreasing chewing rhythmicity. In contrast, low-toughness foods result in jaw movements that are more stereotyped in their timing parameters. In addition, the duration of tooth-food-tooth contact is more variable during the processing of low-stiffness foods compared with tough or stiff foods. Increased toughness is suggested to alter the timing of the movements impacting food fracture whereas increased stiffness may require a more cautious control of jaw movements. This study emphasizes that flexibility in biological movements in response to changes in conditions may not only be observed in timing but also in the variability of their timing within each condition.