Phylogeny, function and ecology in the deep evolutionary history of the mammalian forelimb

Mammals are the only living members of the larger clade Synapsida, which has a fossil record spanning 320 Ma. Despite the fact that much of the ecological diversity of mammals has been considered in the light of limb morphology, the ecological comparability of mammals to their fossil forerunners has not been critically assessed. Because of the wide use of limb morphology in testing ecomorphological hypothesis about extinct tetrapods, we sought: (i) to estimate when in synapsid history, modern mammals become analogues for predicting fossil ecologies; (ii) to document examples of ecomorphological convergence; and (iii) to compare the functional solutions of distinct synapsid radiations. We quantitatively compared the forelimb shapes of the multiple fossil synapsid radiations to a broad sample of extant Mammalia representing a variety of divergent locomotor ecologies. Our results indicate that each synapsid radiation explored different areas of morphospace and arrived at functional solutions that reflected their distinctive ancestral morphologies. This work counters the narrative of non-mammalian synapsid forelimb evolution as a linear progression towards more mammalian morphologies. Instead, a disparate array of early-evolving shapes subsequently contracted towards more mammal-like forms.

Modeling the Determinants of Mechanical Advantage During Jumping: Consequences for Spring- and Muscle-Driven Movement

Systems powered by elastic recoil need a latch to prevent motion while a spring is loaded but allow motion during spring recoil. Some jumping animals that rely on elastic recoil use the increasing mechanical advantage of limb extensor muscles to accomplish latching. We examined the ways in which limb morphology affects latching and the resulting performance of an elastic-recoil mechanism. Additionally, because increasing mechanical advantage is a consequence of limb extension that may be found in many systems, we examined the mechanical consequences for muscle in the absence of elastic elements. By simulating muscle contractions against a simplified model of an extending limb, we found that increasing mechanical advantage can limit the work done by muscle by accelerating muscle shortening during limb extension. The inclusion of a series elastic element dramatically improves mechanical output by allowing for additional muscle work that is stored and released from the spring. This suggests that elastic recoil may be beneficial for more animals than expected when assuming peak isotonic power output from muscle during jumping. The mechanical output of elastic recoil depends on limb morphology; long limbs moving small loads maximize total work, but it is done at a low power, whereas shorter limbs moving larger loads do less work at a higher power. This work-power trade-off of limb morphology is true with or without an elastic element. Systems with relatively short limbs may have performance that is robust to variable conditions such as body mass or muscle activation, while long-limbed systems risk complete failure with relatively minor perturbations. Finally, a changing mechanical advantage latch allows for muscle work to be done simultaneously with spring recoil, changing the predictions for spring mechanical properties. Overall, the design constraints revealed by considering the mechanics of this particular latch will inform our understanding of the evolution of elastic-recoil mechanisms and our attempts to engineer similar systems.

Expression of a hindlimb-determining factor Pitx1 in the forelimb of the lizard Pogona vitticeps during morphogenesis

With over 9000 species, squamates, which include lizards and snakes, are the largest group of reptiles and second-largest order of vertebrates, spanning a vast array of appendicular skeletal morphology. As such, they provide a promising system for examining developmental and molecular processes underlying limb morphology. Using the central bearded dragon ( Pogona vitticeps ) as the primary study model, we examined limb morphometry throughout embryonic development and characterized the expression of three known developmental genes ( GHR, Pitx1 and Shh ) from early embryonic stage through to hatchling stage via reverse transcription quantitative polymerase chain reaction (RT-qPCR) and immunohistochemistry (IHC). In this study, all genes were found to be transcribed in both the forelimbs and hindlimbs of P. vitticeps. While the highest level of GHR expression occurred at the hatchling stage, Pitx1 and Shh expression was greatest earlier during embryogenesis, which coincides with the onset of the differentiation between forelimb and hindlimb length. We compared our finding of Pitx1 expression—a hindlimb-determining gene—in the forelimbs of P. vitticeps to that in a closely related Australian agamid lizard, Ctenophorus pictus , where we found Pitx1 expression to be more highly expressed in the hindlimb compared with the forelimb during early and late morphogenesis—a result consistent with that found across other tetrapods. Expression of Pitx1 in forelimbs has only rarely been documented, including via in situ hybridization in a chicken and a frog. Our findings from both RT-qPCR and IHC indicate that further research across a wider range of tetrapods is needed to more fully understand evolutionary variation in molecular processes underlying limb morphology.

The relationships of the Euparkeriidae and the rise of Archosauria

For the first time, a phylogenetic analysis including all putative euparkeriid taxa is conducted, using a large data matrix analysed with maximum parsimony and Bayesian analysis. Using parsimony, the putative euparkeriid Dorosuchus neoetus from Russia is the sister taxon to Archosauria + Phytosauria. Euparkeria capensis is placed one node further from the crown, and forms a euparkeriid clade with the Chinese taxa Halazhaisuchus qiaoensis and ‘ Turfanosuchus shageduensis ’ and the Polish taxon Osmolskina czatkowicensis . Using Bayesian methods, Osmolskina and Halazhaisuchus are sister taxa within Euparkeriidae, in turn sister to ‘ Turfanosuchus shageduensis ’ and then Euparkeria capensis . Dorosuchus is placed in a polytomy with Euparkeriidae and Archosauria + Phytosauria. Although conclusions remain tentative owing to low node support and incompleteness, a broad phylogenetic position close to the base of Archosauria is confirmed for all putative euparkeriids, and the ancestor of Archosauria +Phytosauria is optimized as similar to euparkeriids in its morphology. Ecomorphological characters and traits are optimized onto the maximum parsimony strict consensus phylogeny presented using squared change parsimony. This optimization indicates that the ancestral archosaur was probably similar in many respects to euparkeriids, being relatively small, terrestrial, carnivorous and showing relatively cursorial limb morphology; this Bauplan may have underlain the exceptional radiaton and success of crown Archosauria.