The ecomorphology of facultative bipedality in Lepidosauria: implications for the evolution of reptilian bipedality

Bipedality is a distinctive locomotor characteristic of some of the most noteworthy animals of all time, including dinosaurs and humans. However, the evolution of a bipedal locomotor mode is poorly understood in reptiles. It has been repeatedly hypothesised that a facultative locomotor mode, where an animal moves both bipedally and quadrupedally under different conditions, forms an intermediate stage in the evolution of obligate bipedality. I demonstrate that the evidence supporting this hypothesis is lacking, recovering facultative bipedality as an intermediate stage only once in multiple independent evolutions of bipedality, under two different topologies. In order to better understand facultative bipedality and the associated anatomies, I performed multiple studies into the ecomorphology and evolution of this behaviour in a modern clade: Lepidosauria. Linear morphometric studies accounting for variation in body size indicate that forelimb segment lengths across locomotor modes do not differ for lepidosaurs of the same size, but that distal hindlimbs segments differ greatly, contrasting with historical tropes. Using 3D landmark-based geometric morphometrics, I demonstrate that arboreal and facultatively bipedal species share many characteristics in the bony elements of the pelvis, including a straight-to-concave iliac blade and large ischial base. These shared anatomies are functionally qualified based on anatomical studies of lepidosaur pelvic girdle myology, and indicate a similarity in mechanical demands of both arboreality and facultative bipedality. Finally, I tested for the correlated evolution of facultative bipedality with substrate preference in a derived clade of squamates: Episquamata. Findings suggest that there is no correlation between substrate and a facultative locomotor mode, instead indicating that facultative bipedality is an exaptation of anatomies associated with vertically diverse environments. This echoes the evolution facultative bipedality in hominin, macropods and rodents, and is distinct from any current hypotheses concerning the evolution of archosaurian bipedality

The relationship between body shape, body size and locomotor mode in extant lepidosaurs.

Despite historic work, the mechanisms and evolutionary drivers associated with the adoption of a facultatively bipedal locomotor mode in extant lepidosaurs are unclear. Recent work has provided insights into the biomechanical triggers of bipedal locomotion, but the associated anatomies are yet to be fully understood, particularly with regard to body size across Lepidosauria. Using a dataset derived from museum specimens, representing a range of lepidosaur body shapes, we highlight the differences between obligate quadrupeds and facultative bipeds within this group and demonstrate the value of non-caudal skeletal material in identifying facultative bipeds using osteology alone. We use multiple statistical approaches to identify trends across locomotor modes relative to body size. Body size has a significant effect upon body proportions across the two locomotor modes, especially in the hindlimbs. Forelimbs lengths do not differ significantly across locomotor modes for animals of similar body size, but distal hindlimbs are significantly longer in facultative bipeds. Interestingly, femoral length does not differ across locomotor modes of a similar body size. Our findings contrast with historical tropes, and are significant for future work attempting to identify the factors driving the evolution of a facultatively bipedal locomotor mode in Lepidosauria

Testing for a facultative locomotor mode in the acquisition of archosaur bipedality

Bipedal locomotion is a defining characteristic of humans and birds and has a profound effect on how these groups interact with their environment. Results from extensive hominin research indicate that there exists an intermediate stage in hominin evolution—facultative bipedality—between obligate quadrupedality and obligate bipedality that uses both forms of locomotion. It is assumed that archosaur locomotor evolution followed this sequence of functional and hence character-state evolution. However, this assumption has never been tested in a broad phylogenetic context. We test whether facultative bipedality is a transitionary state of locomotor mode evolution in the most recent early archosaur phylogenies using maximum-likelihood ancestral state reconstructions for the first time. Across a total of seven independent transitions from quadrupedality to a state of obligate bipedality, we find that facultative bipedality exists as an intermediary mode only once, despite being acquired a total of 14 times. We also report more independent acquisitions of obligate bipedality in archosaurs than previously hypothesized, suggesting that locomotor mode is more evolutionarily fluid than expected and more readily experimented with in these reptiles

The pelvis as an anatomical indicator for facultative bipedality and substrate use in lepidosaurs

Facultative bipedality is regarded as an enigmatic middle ground in the evolution of obligate bipedality and is associated with high mechanical demands in extant lepidosaurs. Traits linked with this phenomenon are largely associated with the caudal end of the animal: hindlimbs and tail. The articulation of the pelvis with both of these structures suggests a morphofunctional role in the use of a facultative locomotor mode. Using a three-dimensional geometric morphometric approach, we examine the pelvic osteology and associated functional implications for 34 species of extant lepidosaur. Anatomical trends associated with the use of a bipedal locomotor mode and substrate preferences are correlated and functionally interpreted based on musculoskeletal descriptions. Changes in pelvic osteology associated with a facultatively bipedal locomotor mode are similar to those observed in species preferring arboreal substrates, indicating shared functionality between these ecologies

The pelvis as an anatomical indicator for facultative bipedality and substrate use in lepidosaurs

AbstractFacultative bipedality is regarded as an enigmatic middle ground in the evolution of obligate bipedality and is associated with high mechanical demands in extant lepidosaurs. Traits linked with this phenomenon are largely associated with the caudal end of the animal: hindlimbs and tail. The articulation of the pelvis with both of these structures suggests a morphofunctional role in the use of a facultative locomotor mode. Using a three-dimensional geometric morphometric approach, we examine the pelvic osteology and associated functional implications for 34 species of extant lepidosaur. Anatomical trends associated with the use of a bipedal locomotor mode and substrate preferences are correlated and functionally interpreted based on musculoskeletal descriptions. Changes in pelvic osteology associated with a facultatively bipedal locomotor mode are similar to those observed in species preferring arboreal substrates, indicating shared functionality between these ecologies.</jats:p

Table-S1-locomotor-diagnoses LG CvB DN supporting 'Testing for a facultative locomotor mode in the acquisition of archosaur bipedality'

The uploaded file consists of a simple data table to support the associated article, "Testing for a facultative locomotor mode in the acquisition of archosaur bipedality". The table outlines the results of a literature survey of all taxa included in the two independent character matrices discussed in the article. The table records the most recent interpretation of locomotor mode for each taxon. The literature survey was conducted to determine whether each taxon was classified as an obligate quadruped (OQ), facultative biped (FB), or obligate biped (OB)

Testing for a facultative locomotor mode in the acquisition of archosaur bipedality.

Bipedal locomotion is a defining characteristic of humans and birds and has a profound effect on how these groups interact with their environment. Results from extensive hominin research indicate that there exists an intermediate stage in hominin evolution-facultative bipedality-between obligate quadrupedality and obligate bipedality that uses both forms of locomotion. It is assumed that archosaur locomotor evolution followed this sequence of functional and hence character-state evolution. However, this assumption has never been tested in a broad phylogenetic context. We test whether facultative bipedality is a transitionary state of locomotor mode evolution in the most recent early archosaur phylogenies using maximum-likelihood ancestral state reconstructions for the first time. Across a total of seven independent transitions from quadrupedality to a state of obligate bipedality, we find that facultative bipedality exists as an intermediary mode only once, despite being acquired a total of 14 times. We also report more independent acquisitions of obligate bipedality in archosaurs than previously hypothesized, suggesting that locomotor mode is more evolutionarily fluid than expected and more readily experimented with in these reptiles.NERC DTP 177291