J. Keleher

Mechanical challenges to freshwater residency in sharks and rays

ABSTRACT Major transitions between marine and freshwater habitats are relatively infrequent, primarily as a result of major physiological and ecological challenges. Few species of cartilaginous fish have evolved to occupy freshwater habitats. Current thought suggests that the metabolic physiology of sharks has remained a barrier to the diversification of this taxon in freshwater ecosystems. Here, we demonstrate that the physical properties of water provide an additional constraint for this species-rich group to occupy freshwater systems. Using hydromechanical modeling, we show that occurrence in fresh water results in a two- to three-fold increase in negative buoyancy for sharks and rays. This carries the energetic cost of lift production and results in increased buoyancy-dependent mechanical power requirements for swimming and increased optimal swim speeds. The primary source of buoyancy, the lipid-rich liver, offers only limited compensation for increased negative buoyancy as a result of decreasing water density; maintaining the same submerged weight would involve increasing the liver volume by very large amounts: 3- to 4-fold in scenarios where liver density is also reduced to currently observed minimal levels and 8-fold without any changes in liver density. The first data on body density from two species of elasmobranch occurring in freshwater (the bull shark Carcharhinus leucas, Müller and Henle 1839, and the largetooth sawfish Pristis pristis, Linnaeus 1758) support this hypothesis, showing similar liver sizes as marine forms but lower liver densities, but the greatest negative buoyancies of any elasmobranch studied to date. Our data suggest that the mechanical challenges associated with buoyancy control may have hampered the invasion of freshwater habitats in elasmobranchs, highlighting an additional key factor that may govern the predisposition of marine organisms to successfully establish in freshwater habitats. Summary: The high negative buoyancy of elasmobranchs increases the cost of locomotion and may be responsible for the scarcity of sharks in fresh water.

Are vertical migrations driven by circadian behaviour? Decoupling of activity and depth use in a large riverine elasmobranch, the freshwater sawfish (Pristis pristis)

Circadian rhythms occur widely amongst living organisms, often in response to diel changes in environmental conditions. In aquatic animals, circadian activity is often synchronised with diel changes in the depths individuals occupy and may be related to predator–prey interactions, where the circadian rhythm is determined by ambient light levels, or have a thermoregulatory purpose, where the circadian rhythm is governed by temperature. Here, these two hypotheses are examined using animal-attached accelerometers in juvenile freshwater sawfish occupying a riverine environment displaying seasonal changes in thermal stratification. Across seasons, diel patterns of depth use (shallow at night and deep in the day) tended to occur only in the late dry seasons when the water was stratified, whereas individuals were primarily shallow in the early dry season which featured no thermal stratification. Activity was elevated during crepuscular and nocturnal periods compared to daytime, regardless of the thermal environment. Our observation of resting at cooler depths is consistent with behavioural thermoregulation to reduce energy expenditure, whereas activity appears linked to ambient light levels and predator–prey interactions. This suggests that circadian rhythms in activity and vertical migrations are decoupled in this species and respond to independent environmental drivers.