Ryan D. Day

Pectoral fin coordination and gait transitions in steadily swimming juvenile reef fishes

SUMMARY A common feature of animal locomotion is its organization into gaits with distinct patterns of movement and propulsor use for specific speeds. In terrestrial vertebrates, limb gaits have been extensively studied in diverse taxa and gait transitions have been shown to provide efficient locomotion across a wide range of speeds. In contrast, examination of gaits in fishes has focused on axial gaits and the transition between synchronous paired fin locomotion and axial propulsion. Because many fishes use their pectoral fins as their primary propulsors, we aimed to examine more broadly the use of pectoral fin gaits in locomotion. We used juvenile reef fishes in these experiments because their swimming could be recorded readily across a wide range of Reynolds numbers, which we thought would promote gait diversity. Based on previous work in larval fishes, we hypothesized that juveniles have alternating pectoral fin movements rather than the synchronous, or in-phase, coordination pattern of adults. In flow tank swim studies, we found that juvenile sapphire damselfish Pomacentrus pavo used two fin gaits during steady swimming. Below approximately 3 BL s-1, P. pavo primarily swam with alternating fin strokes 180° out of phase with one another. At speeds in the range of 3-4 BL s-1, they performed a gait transition to synchronous fin coordination. Between approximately 4 and 8 BL s-1, P. pavo primarily beat their fins synchronously. At around 8 BL s-1 there was another gait transition to body-caudal fin swimming, in which the pectoral fins were tucked against the body. We suggest that the transition from alternating to synchronous fin coordination occurs due to mechanical limits of gait performance rather than to energy efficiency, stability or transitions in hydrodynamic regime. To determine whether this gait transition was species-specific, we surveyed pectoral fin locomotion in juveniles from 11 species in three reef fish families (Pomacentridae, Labridae and Scaridae). We found that this gait transition occurred in every species examined, suggesting that it may be a common behavior of juvenile reef fishes. Greater inclusion of early life history stages in the study of fin-based locomotion should significantly enhance and inform the growing body of work on these behaviors.

Widely used marine seismic survey air gun operations negatively impact zooplankton

Zooplankton underpin the health and productivity of global marine ecosystems. Here we present evidence that suggests seismic surveys cause significant mortality to zooplankton populations. Seismic surveys are used extensively to explore for petroleum resources using intense, low-frequency, acoustic impulse signals. Experimental air gun signal exposure decreased zooplankton abundance when compared with controls, as measured by sonar (~3–4 dB drop within 15–30 min) and net tows (median 64% decrease within 1 h), and caused a two- to threefold increase in dead adult and larval zooplankton. Impacts were observed out to the maximum 1.2 km range sampled, which was more than two orders of magnitude greater than the previously assumed impact range of 10 m. Although no adult krill were present, all larval krill were killed after air gun passage. There is a significant and unacknowledged potential for ocean ecosystem function and productivity to be negatively impacted by present seismic technology.

Why can't young fish eat plants? Neither digestive enzymes nor gut development preclude herbivory in the young of a stomachless marine herbivorous fish

Most young fishes lack the ability to function as herbivores, which has been attributed to two aspects of the digestive system: elevated nitrogen demand and a critical gut capacity. We compared the digestive morphology and biochemistry of two size classes of the marine herbivore Hyporhamphus regularis ardelio, pre-ontogenetic trophic shift (pre-OTS, <100mm) and post-ontogenetic trophic shift (post-OTS, >100mm), to determine what limits the onset of herbivory and how their digestive processes fit with current models of digestion. Two gut-somatic indices comparing gut length to body length (relative gut length) and body mass (Zihlers Index) demonstrated a significant decrease (RGL 0.59→0.49, P<0.01; ZI 3.24→2.44, P<0.01) in gut length relative to body size. There was little difference in enzyme activity between the two classes, with juveniles showing similar levels of carbohydrase and lipase and less protease compared with adults, indicating that juveniles did not preferentially target nitrogen and were as capable of digesting an herbivorous diet. These findings suggest that herbivory in this fish is not limited by the function of the post-oesophageal digestive tract, but rather the ability of the pharyngeal mill to mechanically process plants. Our findings offer partial support for the current model of stomachless digestion, indicating that further refinement may be necessary.

Functional morphology of digestion in the stomachless, piscivorous needlefishes Tylosurus gavialoides and Strongylura leiura ferox (Teleostei: Beloniformes).

Belonidae are unusual in that they are carnivorous but lack a stomach and have a straight, short gut. To develop a functional morphological model for this unusual system the gut contents and alimentary tract morphology of Tylosurus gavialoides and Strongylura leiura ferox were investigated. The posterior orientation of the majority of the pharyngeal teeth supports the swallowing of whole large prey, but not their mastication. Mucogenic cells are abundant in the mucosa lining, particularly the esophagus, and their secretions are likely to protect the gut lining from damage while lubricating passage of the prey. Esophagus, anterior intestine, posterior intestine, and rectum all have highly reticulate mucosae. The anterior three gut sections are distensible to accommodate the passage of prey. However, following ingestion large prey are passed to the highly distensible posterior intestine where they rest head first against the ileorectal valve. Alimentary pH ranges from neutral to weakly acidic. Fish prey is digested head first with the head being largely digested while the remainder of the body is still intact. The nondistensibility of the rectum and the small aperture provided by the ileorectal valve suggest the products of intestinal digestion are either small particulates or fluids that pass into rectum where they are absorbed. J. Morphol. 2009.

Seismic air gun exposure during early-stage embryonic development does not negatively affect spiny lobster Jasus edwardsii larvae (Decapoda:Palinuridae)

Marine seismic surveys are used to explore for sub-seafloor oil and gas deposits. These surveys are conducted using air guns, which release compressed air to create intense sound impulses, which are repeated around every 8–12 seconds and can travel large distances in the water column. Considering the ubiquitous worldwide distribution of seismic surveys, the potential impact of exposure on marine invertebrates is poorly understood. In this study, egg-bearing female spiny lobsters (Jasus edwardsii) were exposed to signals from three air gun configurations, all of which exceeded sound exposure levels (SEL) of 185 dB re 1 μPa2·s. Lobsters were maintained until their eggs hatched and the larvae were then counted for fecundity, assessed for abnormal morphology using measurements of larval length and width, tested for larval competency using an established activity test and measured for energy content. Overall there were no differences in the quantity or quality of hatched larvae, indicating that the condition and development of spiny lobster embryos were not adversely affected by air gun exposure. These results suggest that embryonic spiny lobster are resilient to air gun signals and highlight the caution necessary in extrapolating results from the laboratory to real world scenarios or across life history stages.

Development of the pharyngeal dentition of two herbivorous halfbeaks (Teleostei : Hemiramphidae) and implications for the hemiramphid ontogenetic trophic shift

Development of the pharyngeal dentition of two herbivorous halfbeaks, Hyporhamphus regularis ardelio (Whitley, 1931) and Arrhamphus sclerolepis krefftii (Steindachner, 1867), was examined quantitatively to assess features that might confer their ability to shift their diet from animal to plant material. Toothed area, tooth number, maximum tooth diameter and tooth wear area in both pharyngeal tooth pads of both taxa increased with ontogeny, whereas tooth density decreased. Comparing individuals of the two taxa at similar standard lengths indicated that A. sclerolepis krefftii showed hypertrophy of the majority of pharyngeal characters in relation to H. regularis ardelio of a similar standard length. That A. sclerolepis krefftii is more developmentally advanced than H. regularis ardelio in almost all dentigerous characters studied indicates that pharyngeal development may allow the former to commence herbivory at a smaller standard length than the latter species. The evolutionary and ecological implications of these findings are discussed in the context of a group of fishes that is overexploited worldwide.

Ballistic Beloniformes attacking through Snell's Window.

Needlefishes (Beloniformes) were observed employing a range of stalking and attacking behaviours to attack schools of bait fishes ranging from the use of tactics common to predatory fishes to a novel behaviour: the use of leaping, aerial attacks. These aerial attacks are suggested to serve two purposes: to extend the attack range of the needlefishes and to reduce their preys potential for evasion. Furthermore, a third purpose is hypothesized that the needlefishes are taking advantage of Snells Window, an optical effect which may mask their approach to their prey.

Ultrastructure and the importance of wear in the dentition of the halfbeak (Pisces: Hemiramphidae) pharyngeal mill

To assess how tooth microstructure and composition might facilitate the pharyngeal mill mechanism of halfbeaks, apatite structure and iron content were determined by scanning electron microscopy and energy dispersive X‐ray analysis for Hyporhamphus regularis ardelio, Arrhamphus sclerolepis krefftii, and Hemiramphus robustus. Iron was present in developing teeth and was concentrated along the shearing edge of spatulate incisiform teeth, which dominate the occlusive wear zone in all three species. A model based on tooth structure and wear rate is proposed to explain how halfbeaks maintain a fully functional occlusion zone throughout growth and consequent tooth addition and replacement. Replacement teeth erupt and wear rapidly so that a constant occlusion plane is always present. Iron within the tooth tissue reduces the wear rate of the cutting edge while simultaneously maintaining its sharpness and efficiency. J. Morphol. 2009.