They like to move it (move it): walking kinematics of balitorid loaches of Thailand.

Balitorid loaches are a family of fishes that exhibit morphological adaptations to living in fast flowing water, including an enlarged sacral rib that creates a 'hip'-like skeletal connection between the pelvis and the axial skeleton. The presence of this sacral rib, the robustness of which varies across the family, is hypothesized to facilitate terrestrial locomotion seen in the family. Terrestrial locomotion in balitorids is unlike that of any known fish: the locomotion resembles that of terrestrial tetrapods. Emergence and convergence of terrestrial locomotion from water to land has been studied in fossils; however, studying balitorid walking provides a present-day natural laboratory to examine the convergent evolution of walking movements. We tested the hypothesis that balitorid species with more robust connections between the pelvic and axial skeleton (M3 morphotype) are more effective at walking than species with reduced connectivity (M1 morphotype). We predicted that robust connections would facilitate travel per step and increase mass support during movement. We collected high-speed video of walking in seven balitorid species to analyze kinematic variables. The connection between internal anatomy and locomotion on land are revealed herein with digitized video analysis, µCT scans, and in the context of the phylogenetic history of this family of fishes. Our species sampling covered the extremes of previously identified sacral rib morphotypes, M1 and M3. Although we hypothesized the robustness of the sacral rib to have a strong influence on walking performance, there was not a large reduction in walking ability in the species with the least modified rib (M1). Instead, walking kinematics varied between the two balitorid subfamilies with a generally more 'walk-like' behavior in the Balitorinae and more 'swim-like' behavior in the Homalopteroidinae. The type of terrestrial locomotion displayed in balitorids is unique among living fishes and aids in our understanding of the extent to which a sacral connection facilitates terrestrial walking.

Sucker with a fat lip: The soft tissues underlying the viscoelastic grip of remora adhesion.

Remoras are fishes that attach to a broad range of hosts using an adhesive disc on their head that is derived from dorsal fin elements. Research on the adhesive mechanism of remoras has focused primarily on the skeletal components of the disc and their contribution to generating suction and friction. However, the soft tissues of the disc, such as the soft lip surrounding the bony disc and the muscles that control the bony lamellae, have been largely ignored. To understand the sealing mechanism of the disc, it is imperative to understand the tissue morphology and material properties of the soft lip. Here, we show that the soft lip surrounding the remora disc is comprised of discrete multilayered collagen, fat, and elastic tissues which we hypothesize to have specific roles in the viscoelastic sealing mechanism of the remora disc. The central, heavily vascularized fat and collagen layer are infiltrated by strands of elastic tissue and surrounded by crossed-fiber collagen. A newly described jubilee muscle underneath the adhesive disc provides a mechanism for stopping venous return from the disc lip, thereby allowing it to become engorged and create a pressurized fit to the attachment substrate. Thus, the remora lip acts as a vascular hydrostat.

Channeling vorticity: modeling the filter-feeding mechanism in silver carp using µCT and 3D PIV.

Invasive silver carp are thriving within eutrophic environments in the United States, in part because of their highly efficient filter-feeding mechanism. Silver carp utilize modified gill rakers to capture a specific range of food; however, their greatly modified filtering morphology allows them to feed on phytoplankton and zooplankton ranging in size from 4 to 85 µm. The filtering apparatus of silver carp comprises rigid filtering plates where the outer anatomy of these plates is characterized by long parallel channels that change in orientation along the length of the plate. Here, we investigate the underlying morphology and concomitant hydrodynamics that support the filtration mechanisms of silver and bighead carp. Bighead carp are also invasive filter feeders, but their filtering apparatus is morphologically distinct from that of silver carp. Using 3D particle image velocimetry, we determined how particles and fluid interact with the surface of the gill rakers/plates. Filtering plates in silver carp induce strong directed vortical flow, whereas the filtering apparatus of bighead carp resulted in a type of haphazard cross-flow filtration. The organized vortical flow established by silver carp likely increased the number of interactions that the particle-filled water had with the filtering membrane. This strong vortical organization is maintained only at 0.75 body lengths s-1, and vortical flow is poorly developed and maintained at slower and faster speeds. Moreover, we found that absolute vorticity magnitude in silver carp is an order of magnitude greater than in bighead carp.

Intraspecific variation in feeding and locomotor kinematics during prey capture in redbreast sunfish (Lepomis auritus).

Biomechanics research often revolves around understanding traits impacting suction feeding performance in fishes, using freshwater ray-finned sunfishes (Family Centrarchidae) as models. However, simultaneous feeding and locomotion kinematics during prey capture are not recorded for many species and there is less information on how these kinematics vary within a species and within individuals. To (1) add to existing data on the prey capture kinematics of centrarchids, (2) assess variation in a species both within and across individuals, and (3) compare morphology and prey capture kinematics of well-sampled centrarchids, we filmed five redbreast sunfish (Lepomis auritus) at 500 fps-1 approaching and striking non-evasive prey. Redbreast approach prey at ~30 cm s-1 and use approximately 70% of their maximum gape size. Traits related to feeding are more repeatable than traits related to locomotion. However, the Accuracy Index (AI) was consistent across individuals (AI = 0.76 ± 0.07). Functionally, redbreast sunfish are more similar to bluegill sunfish but morphologically they fall in the intermediate morphospace alongside green sunfish when compared with other centrarchids. These data show that whole organism outcomes (AI) are similar despite variation present both within and across individuals and demonstrate the importance of considering both interspecific and intraspecific differences in the functional diversity of ecologically and evolutionarily important behaviors such as prey capture.

It Pays to Be Bumpy: Drag Reducing Armor in the Pacific Spiny Lumpsucker, Eumicrotremus orbis.

Armor is a multipurpose set of structures that has evolved independently at least 30 times in fishes. In addition to providing protection, armor can manipulate flow, increase camouflage, and be sexually dimorphic. There are potential tradeoffs in armor function: increased impact resistance may come at the cost of maneuvering ability; and ornate armor may offer visual or protective advantages, but could incur excess drag. Pacific spiny lumpsuckers (Eumicrotremus orbis) are covered in rows of odontic, cone-shaped armor whorls, protecting the fish from wave driven impacts and the threat of predation. We are interested in measuring the effects of lumpsucker armor on the hydrodynamic forces on the fish. Bigger lumpsuckers have larger and more complex armor, which may incur a greater hydrodynamic cost. In addition to their protective armor, lumpsuckers have evolved a ventral adhesive disc, allowing them to remain stationary in their environment. We hypothesize a tradeoff between the armor and adhesion: little fish prioritize suction, while big fish prioritize protection. Using micro-CT, we compared armor volume to disc area over lumpsucker development and built 3D models to measure changes in drag over ontogeny. We found that drag and drag coefficients decrease with greater armor coverage and vary consistently with orientation. Adhesive disc area is isometric but safety factor increases with size, allowing larger fish to remain attached in higher flows than smaller fish.

Knowing when to stick: touch receptors found in the remora adhesive disc.

Remoras are fishes that piggyback onto larger marine fauna via an adhesive disc to increase locomotor efficiency, likelihood of finding mates and access to prey. Attaching rapidly to a large, fast-moving host is no easy task, and while research to date has focused on how the disc supports adhesion, no attention has been paid to how or if remoras are able to sense attachment. We identified push-rod-like mechanoreceptor complexes embedded in the soft lip of the remora adhesive disc that are known in other organisms to respond to touch and shear forces. This is, to our knowledge, the first time such mechanoreceptor complexes are described in fishes as they were only known previously in monotremes. The presence of push-rod-like mechanoreceptor complexes suggests not only that fishes may be able to sense their environment in ways not heretofore described but that specialized tactile mechanoreceptor complexes may be a more basal vertebrate feature than previously thought.

Skeletal and muscular pelvic morphology of hillstream loaches (Cypriniformes: Balitoridae).

The rheophilic hillstream loaches (Balitoridae) of South and Southeast Asia possess a range of pelvic girdle morphologies, which may be attributed to adaptations for locomotion against rapidly flowing water. Specifically, the connectivity of the pelvic plate (basipterygium) to the vertebral column via a sacral rib, and the relative size and shape of the sacral rib, fall within a spectrum of three discrete morphotypes: long, narrow rib that meets the basipterygium; thicker, slightly curved rib meeting the basipterygium; and robust crested rib interlocking with the basipterygium. Species in this third category with more robust sacral rib connections between the basipterygium and vertebral column are capable of walking out of water with a tetrapod-like lateral-sequence, diagonal-couplet gait. This behavior has not been observed in species lacking direct skeletal connection between the vertebrae and the pelvis. The phylogenetic positions of the morphotypes were visualized by matching the morphological features onto a novel hypothesis of relationships for the family Balitoridae. The morphotypes determined through skeletal morphology were correlated with patterns observed in the pelvic muscle morphology of these fishes. Transitions towards increasingly robust pelvic girdle attachment were coincident with a more anterior origin on the basipterygium and more lateral insertion of the muscles on the fin rays, along with a reduction of the superficial abductors and adductors with more posterior insertions. These modifications are expected to provide a mechanical advantage for generating force against the ground. Inclusion of the enigmatic cave-adapted balitorid Cryptotora thamicola into the most data-rich balitorid phylogeny reveals its closest relatives, providing insight into the origin of the skeletal connection between the axial skeleton and basipterygium.

Complete mitochondrial genome of the winghead shark, Eusphyra blochii (Elasmobranchii: Sphyrnidae).

The complete mitochondrial genome (16,726 bp) of the winghead shark, Eusphyra blochii is presented. This species is exploited throughout parts of its range, and is currently listed as Near Threatened by the IUCN Red List. A phylogenetic analysis placed E. blochii within the Carcharhiniformes, as a sister taxon to Sphyrna lewini and S. zygaena.

Material Properties of Hagfish Skin, with Insights into Knotting Behaviors.

Hagfishes (Myxinidae) often integrate whole-body knotting movements with jawless biting motions when reducing large marine carcasses to ingestible items. Adaptations for these behaviors include complex arrangements of axial muscles and flexible, elongate bodies without vertebrae. Between the axial muscles and the hagfish skin is a large, blood-filled subcutaneous sinus devoid of the intricate, myoseptal tendon networks characteristic of the taut skins of other fishes. We propose that the loose-fitting skin of the hagfish facilitates the formation and manipulation of body knots, even if it is of little functional significance to steady swimming. Hagfish skin is a relatively thick, anisotropic, multilayered composite material comprising a superficial, thin, and slimy epidermis, a middle dermal layer densely packed with fibrous tissues, and a deep subdermal layer comprised of adipose tissue. Hagfish skin is stiffer when pulled longitudinally than circumferentially. Stress-strain data from uniaxial tensile tests show that hagfish skins are comparable in tensile strength and stiffness to the taut skins of elongate fishes that do not engage in knotting behaviors (e.g., sea lamprey and penpoint gunnel). Sheath-core-constructed ropes, which serve as more accurate models for hagfish bodies, demonstrate that loose skin (extra sheathing) enhances flexibility of the body (rope). Along with a loose-fitting skin, the morphologies of hagfish skin parallel those of moray eels, which are also known for generating and manipulating figure-eight-style body knots when struggling with prey.