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Synopsis Animals clinging to natural surfaces have to generate attachment across a range of surface roughnesses in both dry and wet conditions. Plethodontid salamanders can be aquatic, semi-aquatic, terrestrial, arboreal, troglodytic, saxicolous, and fossorial and therefore may need to climb on and over rocks, tree trunks, plant leaves, and stems, as well as move through soil and water. Sixteen species of salamanders were tested to determine the effects of substrate roughness and wetness on maximum cling angle. Substrate roughness had a significant effect on maximum cling angle, an effect that varied among species. Substrates of intermediate roughness (asperity size 100–350 µm) resulted in the poorest attachment performance for all species. Small species performed best on smooth substrates, while large species showed significant improvement on the roughest substrates (asperity size 1000–4000 µm), possibly switching from mucus adhesion on a smooth substrate to an interlocking attachment on rough substrates. Water, in the form of a misted substrate coating and a flowing stream, decreased cling performance in salamanders on smooth substrates. However, small salamanders significantly increased maximum cling angle on wetted substrates of intermediate roughness, compared with the dry condition. Study of cling performance and its relationship to surface properties may cast light onto how this group of salamanders has radiated into the most speciose family of salamanders that occupies diverse habitats across an enormous geographical range.

The evolution of ballistic tongue projection in plethodontid salamanders—a high-performance and thermally robust musculoskeletal system—is ideal for examining how the components required for extreme performance in animal movement are assembled in evolution. Our comparative data on whole-organism performance measured across a range of temperatures and the musculoskeletal morphology of the tongue apparatus were examined in a phylogenetic framework and combined with data on muscle contractile physiology and neural control. Our analysis reveals that relatively minor evolutionary changes in morphology and neural control have transformed a muscle-powered system with modest performance and high thermal sensitivity into a spring-powered system with extreme performance and functional robustness in the face of evolutionarily conserved muscle contractile physiology. Furthermore, these changes have occurred in parallel in both major clades of this largest family of salamanders. We also find that high-performance tongue projection that exceeds available muscle power and thermal robustness of performance coevolve, both being emergent properties of the same elastic-recoil mechanism. Among the taxa examined, we find muscle-powered and fully fledged elastic systems with enormous performance differences, but no intermediate forms, suggesting that incipient elastic mechanisms do not persist in evolutionary time. A growing body of data from other elastic systems suggests that similar coevolution of traits may be found in other ectothermic animals with high performance, particularly those for which thermoregulation is challenging or ecologically costly.

Plethodontid salamanders inhabit terrestrial, scansorial, arboreal, and troglodytic habitats in which clinging and climbing allow them access to additional food and shelter as well as escape from unfavorable temperature and moisture conditions and ground-dwelling predators. Although salamanders lack claws and toe pads found on other taxa, they successfully cling to and climb on inclined, vertical, and inverted substrates in nature. Maximum cling angle was tested on smooth acrylic, and the relationship between cling angle, body mass, and surface area of attachment (contact area) was investigated. This study found that many salamander species can cling fully inverted using only a portion of their ventral surface area to attach. Salamanders fall into three functional groups based on mass and maximum cling angle: (1) high performing, very small salamanders, (2) moderately high performing small and medium-sized salamanders, and (3) low performing large salamanders. They show significant differences in maximum cling angle, even between species of similar mass. In species of similar mass experiencing significantly different detachment stress (resulting from significantly different contact area), differences in morphology or behavior affect how much body surface is attached to the substrate. High performance in some species, such as Desmognathus quadramaculatus, is attributable to large contact area; low performance in a similarly sized species, Ensatina eschscholtzii, is due to behavior which negatively impacts contact area. There was not clear evidence of scaling of adhesive strength with increasing body size. Salamander maximum cling angle is the result of morphology and behavior impacting the detachment stresses experienced during clinging.

Black flies are ubiquitous and important members of lotic ecosystems. Size is known to affect many aspects of their life in the aquatic larval stage, including intraspecific competition for feeding sites. As filter feeders, flow affects their ability to feed and reach sufficiently fast flow. This, in turn, can lead to risky fluid-mediated dispersal behavior in search of better conditions. It is surprising, therefore, that little information is available regarding how physiological and environmental factors combine to affect larval growth rates. The present study determines the relative growth rates of small (0.6 mm) and large (approximately 4 mm) larvae in laboratory flumes designed to produce spatially homogeneous and temporally consistent flow regimes at ecologically relevant velocities (44 and 64 cm/s). Our results indicate that size and flow both influence growth rates and that the 2 interact significantly. Young larvae exhibit faster growth rates and a greater positive response to increased flow speed. This result might help explain why smaller larvae have a greater propensity to disperse than larger larvae: the benefit of increased growth rate that they receive from relocating to faster flow might balance the risks inherent in dispersal.

Elastic-recoil mechanisms can improve organismal performance and circumvent the thermal limitations of muscle contraction, yet they require the appropriate motor control to operate. We compare muscle activity during tongue projection in salamanders with elastically powered, ballistic projection with activity of those with muscle-powered, non-ballistic projection across a range of temperatures to understand how motor control is integrated with elastically powered movements, and how this integration contributes to reduced thermal sensitivity. Species with ballistic tongue projection activated and deactivated their projector muscles significantly earlier than non-ballistic species, in a pattern consistent with a mechanism in which the muscle strains elastic tissue that subsequently recoils to power projection. Tongue projection was more thermally robust in ballistic species, but in both ballistic and non-ballistic species the projector muscles were activated earlier and for longer as temperature decreased. The retractor muscles showed a pattern similar to that of the projector muscles, but declined in a similar manner in the two groups. Muscle activity intensity also decreased at low temperatures in both groups, revealing that compensatory muscle activation does not account for the improved thermal robustness in ballistic species. Thus, relatively minor shifts in motor patterns accompanying morphological changes such as increased elastic tissue are sufficient to improve performance and decrease its thermal sensitivity without specialization of muscle contractile physiology.

The experiments described in this paper have two goals. The first goal is to show how students can perform simple but fundamental measurements of objects moving through simple liquids (such as water, oil, or honey). In doing so, students can verify Stokes' law, which governs the motion of spheres through simple liquids, and see how it fails at higher object speeds. Moreover, they can qualitatively study fluid patterns at various object speeds (Reynolds numbers). The second goal is to help students make connections between physics and other sciences. Specifically, the results of these experiments can be used to help students understand the role of fluid motion in determining the shape of an organism, or where it lives. At Saint Josephs University we have developed these experiments as part of a newly developed course in biomechanics where both physics and biology undergraduate students bring their ideas and expertise to enrich a shared learning environment.